Smithsonian Institution

Scholarly Press

Smithsonian Institution

Scholarly Press

smithsonian contributions to botany • number 100

Smithsonian Institution

Scholarly Press

Smithsonian Institution

Scholarly Press

Flora of Guaramacal

(Venezuela):

Monocotyledons

L. J. Dorr
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smithsonian contributions to botany • numb e r 1 0 0

Flora of Guaramacal

(Venezuela):

Monocotyledons

L. J. Dorr

WASHINGTON D.C.

2014
ABSTRACT. Dorr, L. J. Flora of Guaramacal (Venezuela): Monocotyledons. Smithsonian Contributions to Botany, number 100, xiv + 289 pages, 26 figures, 316 maps, 2014.—The present volume is a taxonomic study of the monocotyledons 

that occur in Guaramacal National Park (officially El Parque Nacional “General Cruz Carillo en Guaramacal”), which 

protects the Ramal de Guaramacal, an outlier of the more extensive Cordillera de Mérida in the Venezuelan Andes. The 

park covers 215 km2 in Trujillo and Portuguesa states and includes forest and páramo found between 1,600 and 3,100 m 

elevation. In all, 25 families, 135 genera, and 315 species (316 taxa) are treated; the greatest species diversity occurs in Orchidaceae (147 species), Poaceae (57 species), and Cyperaceae (28 species). Twenty species are currently known only from 

Guaramacal and might be endemic. Four new combinations are made: Cyperus hortensis (Salzm. ex Steud.) Dorr, Acronia 

archidiaconi (Ames) Carnevali & G. A. Romero, Crocodeilanthe gelida (Lindl.) Carnevali & I. Ramírez, and Fernandezia 

schultesii (L. O. Williams) Carnevali & Dorr.

RESUMEN. Dorr, L. J. Flora de Guaramacal (Venezuela): Monocotiledóneas. Smithsonian Contributions to Botany, 

número 100, xiv + 289 páginas, 26 figuras, 316 mapas, 2014.—El presente volumen es un estudio taxonómico de las 

monocotiledóneas que se encuentran en el Parque Nacional Guaramacal (oficialmente El Parque Nacional “General Cruz 

Carrillo en Guaramacal”), que protege el Ramal de Guaramacal, un parte aislada de la más extensa Cordillera de Mérida 

en los Andes venezolanos. El parque ocupa 215 km2 en los estados Trujillo y Portuguesa e incluye bosques y páramos 

que se encuentran entre 1.600 y 3.100 m sobre el nivel del mar. Un total de 25 familias, 135 géneros, y 315 especies (316 

taxones) son aquí tratados; las familias con mayor diversidad de especies son Orchidaceae (147 especies), Poaceae (57 

especies), y Cyperaceae (28 especies). Veinte especies son conocidas sólamente de Guaramacal las cuales podrían ser endémicas. Cuatro nuevas combinaciones son efectuadas: Cyperus hortensis (Salzm. ex Steud.) Dorr, Acronia archidiaconi

(Ames) Carnevali & G. A. Romero, Crocodeilanthe gelida (Lindl.) Carnevali & I. Ramírez, y Fernandezia schultesii (L. O. 

Williams) Carnevali & Dorr.

Cover images (left to right): Figures 23H, 23G, 25A, 25F, 24E, and 25K.

Published by SMITHSONIAN INSTITUTION SCHOLARLY PRESS

P.O. Box 37012, MRC 957, Washington, D.C. 20013-7012

www.scholarlypress.si.edu

Copyright © 2014 Smithsonian Institution

Text, images, and taxonomic accounts contributed solely by Laurence J. Dorr or by Mark T. Strong are in the public 

domain. The rights to all other text and images in this publication, including cover and interior designs, are owned either 

by the Smithsonian Institution, by contributing authors, or by third parties. Fair use of materials is permitted for personal, educational, or noncommercial purposes. Users must cite author and source of content, must not alter or modify 

copyrighted content, and must comply with all other terms or restrictions that may be applicable. Users are responsible 

for securing permission from a rights holder for any other use.

Library of Congress Cataloging-in-Publication Data

Dorr, Laurence J.

Flora of Guaramacal (Venezuela) : monocotyledons / L. J. Dorr.

 pages cm. — (Smithsonian contributions to botany, ISSN 0081-024X ; number 100)

Other title: Monocotyledons

English with abstracts in English and Spanish.

Includes bibliographical references and index.

1. Monocotyledons—Venezuela—Parque Nacional Guaramacal. 2. Botany—Venezuela—Parque Nacional 

Guaramacal. 3. Parque Nacional Guaramacal (Venezuela) I. Title. II. Title: Monocotyledons. III. Series: Smithsonian 

contributions to botany ; no. 100.

QK273.D67 2014

584.0987—dc23

2014013050

ISSN: 0081-024X (print); 1938-2812 (online)

c The paper used in this publication meets the minimum requirements of the American National Standard for Permanence 

of Paper for Printed Library Materials Z39.48–1992.
Contents

LIST OF FIGURES v

LIST OF DISTRIBUTION MAPS vii

INTRODUCTION 1

Abbreviations and Commonly Used Symbols 2

Contributors 2

SYSTEMATIC TREATMENT 3

Key to the Families of Monocotyledons 3

Alstroemeriaceae 5

Amaryllidaceae 8

Araceae 9

Arecaceae 23

Bromeliaceae 30

Burmanniaceae 44

Cannaceae 45

Commelinaceae 46

Costaceae 50

Cyclanthaceae 52

Cyperaceae 55

Dioscoreaceae 84

Eriocaulaceae 90

Heliconiaceae 92

Hypoxidaceae 95

Iridaceae 96

Juncaceae 101

Marantaceae 105

Orchidaceae 106

Poaceae 209

Smilacaceae 249

Tofieldiaceae 252

Xanthorrhoeaceae 254

Xyridaceae 255

Zingiberaceae 256
i v • SMITHSONIAN CONTRIBUTIONS TO BOTANY

LIST OF NEW COMBINATIONS 260

NOTE ADDED IN PROOF 260

ACKNOWLEDGMENTS 265

REFERENCES 267

INDEX OF SCIENTIFIC NAMES 281

INDEX OF COMMON NAMES 289
Figures

1. Bomarea amilcariana Stergios & Dorr 6

2. Anthurium ramoncaracasii Stergios & Dorr 17

3. Aiphanes lindeniana (H. Wendl.) H. Wendl. 25

4. Tradescantia zanonia (L.) Sw. and Dichorisandra hexandra (Aubl.) 

Kuntze ex Hand.-Mazz. 48

5. Carex jamesonii Boott 58

6. Cyperus niger Ruiz & Pav. 64

7. Eleocharis montana (Kunth) Roem. & Schult. 66

8. Fimbristylis complanata (Retz.) Link 69

9. Oreobolus venezuelensis Steyerm. 71

10. Rhynchospora guaramacalensis M. T. Strong 75

11. Uncinia hamata (Sw.) Urb. 85

12. Dioscorea lisae Dorr & Stergios 89

13. Paepalanthus pilosus (Kunth) Kunth 91

14. Heliconia hirsuta L. f. and H. meridensis Klotzsch 94

15. Hypoxis decumbens L. 97

16. Sisyrinchium tinctorium Kunth and Orthrosanthus acorifolius

(Kunth) Ravenna 99

17. Luzula gigantea Desv. and Juncus microcephalus Kunth 103

18. Stromanthe tonckat (Aubl.) Eichler 107

19. Smilax domingensis Willd. 250

20. Harperocallis robustior (Steyerm.) L. M. Campb. & Dorr 253

21. Xyris subulata var. acutifolia Heimerl 257

22. Renealmia thyrsoidea (Ruiz & Pav.) Poepp. & Endl. subsp. thyrsoidea

and Hedychium coronarium J. Koenig 259

23. Bomarea amilcariana Stergios & Dorr, Anthurium amoenum Kunth 

& C. D. Bouché, A. nymphaeifolium K. Koch & C. D. Bouché, 

A. ramoncaracasii Stergios & Dorr, A. smaragdinum G. S. Bunting, 

A. ginesii Croat, Philodendron fraternum Schott, and Xanthosoma 

sagittifolium (L.) Schott 261

24. Wettinia praemorsa (Willd.) Wess. Boer, Puya aristeguietae L. B. Sm., 

Tillandsia fendleri Griseb., T. complanata Benth., P. venezuelana

L. B. Sm., Mezobromelia capituligera (Griseb.) J. R. Grant, and 

Vriesea incurva (Griseb.) Read 262
v i • SMITHSONIAN CONTRIBUTIONS TO BOTANY

25. Canna jaegeriana Urb., Sphaeradenia laucheana (Sander ex Mast.) 

Harling subsp. laucheana, Carex jamesonii Boott, Paepalanthus pilosus

(Kunth) Kunth, Heliconia meridensis Klotzsch, Elleanthus wageneri

(Rchb. f.) Rchb. f., Sisyrinchium tinctorium Kunth, Anathallis 

sclerophylla (Lindl.) Pridgeon & M. W. Chase, Epidendrum secundum

Jacq., Habenaria gollmeri Schltr., Maxillaria triloris E. Morren 263

26. Chusquea fendleri Munro, Andropogon bicornis L., Cortaderia 

hapalotricha (Pilg.) Conert, Prosthechea brachychila (Lindl.) 

W. E. Higgins, Eccremis coarctata (Ruiz & Pav.) Baker, Smilax 

domingensis Willd. 264
Distribution Maps

1. Bomarea amilcariana 7

2. Bomarea edulis 7

3. Bomarea salicifolia 8

4. Eucharis amazonica 9

5. Anthurium amoenum 11

6. Anthurium bernardii 12

7. Anthurium crassinervium 12

8. Anthurium eminens subsp. eminens 13

9. Anthurium gehrigeri 13

10. Anthurium ginesii 14

11. Anthurium julianii 14

12. Anthurium longegeniculatum 15

13. Anthurium nubicola 15

14. Anthurium nymphaeifolium 16

15. Anthurium ramoncaracasii 16

16. Anthurium scandens subsp. scandens 18

17. Anthurium smaragdinum 18

18. Monstera adansonii var. laniata 19

19. Philodendron fraternum 21

20. Philodendron tuerckheimii 21

21. Rhodospatha badilloi 22

22. Xanthosoma sagittifolium 23

23. Aiphanes lindeniana 24

24. Chamaedorea linearis 26

25. Chamaedorea pinnatifrons 27

26. Geonoma lehmannii subsp. lehmannii 28

27. Geonoma orbignyana subsp. orbignyana 28

28. Geonoma undata subsp. undata 29

29. Prestoea acuminata var. acuminata 29

30. Wettinia praemorsa 30

31. Aechmea spectabilis 32

32. Greigia alborosea 32

33. Guzmania mitis 33

34. Guzmania squarrosa 33

35. Mezobromelia capituligera 34
viii • SMITHSONIAN CONTRIBUTIONS TO BOTANY

36. Pitcairnia brevicalycina 35

37. Pitcairnia hitchcockiana 35

38. Pitcairnia maidifolia 36

39. Puya aristeguietae 37

40. Puya venezuelana 37

41. Racinaea spiculosa var. spiculosa 38

42. Racinaea tetrantha var. caribaea 39

43. Racinaea tetrantha var. miniata 40

44. Tillandsia biflora 41

45. Tillandsia compacta 41

46. Tillandsia complanata 42

47. Tillandsia fendleri 42

48. Tillandsia myriantha 43

49. Vriesea incurva 43

50. Gymnosiphon suaveolens 45

51. Canna jaegeriana 46

52. Commelina obliqua 47

53. Dichorisandra hexandra 49

54. Phyodina gracilis 49

55. Tradescantia zanonia 50

56. Costus guanaiensis var. macrostrobilus 51

57. Costus spiralis var. spiralis 52

58. Asplundia moritziana 53

59. Asplundia vagans 54

60. Sphaeradenia laucheana subsp. laucheana 55

61. Carex bonplandii 57

62. Carex jamesonii 59

63. Carex longii 59

64. Carex tachirensis 60

65. Carex tamana 60

66. Cyperus hermaphroditus 62

67. Cyperus hortensis 63

68. Cyperus niger 63

69. Eleocharis flavescens 65

70. Eleocharis montana 67

71. Eleocharis stenocarpa 67

72. Fimbristylis complanata 68

73. Oreobolus venezuelensis 70

74. Rhynchospora globosa var. globosa 73

75. Rhynchospora gollmeri 74

76. Rhynchospora guaramacalensis 74

77. Rhynchospora immensa 76

78. Rhynchospora lechleri 77

79. Rhynchospora locuples 77

80. Rhynchospora macrochaeta 78

81. Rhynchospora polyphylla 79

82. Rhynchospora rugosa 79

83. Rhynchospora ruiziana 80

84. Rhynchospora tuerckheimii 80

85. Rhynchospora sp. A 81

86. Scleria distans 82

87. Scleria tropicalis 83

88. Uncinia hamata 84

89. Dioscorea coriacea 87
NUMBER 100 • i x

90. Dioscorea lehmannii 87

91. Dioscorea lisae 88

92. Dioscorea meridensis 88

93. Paepalanthus pilosus 92

94. Heliconia hirsuta 93

95. Heliconia meridensis 93

96. Heliconia stricta 95

97. Hypoxis decumbens 96

98. Orthrosanthus acorifolius 98

99. Sisyrinchium micranthum 100

100. Sisyrinchium tinctorium 101

101. Juncus bufonius 102

102. Juncus microcephalus 104

103. Luzula gigantea 104

104. Stromanthe jacquinii 106

105. Stromanthe tonckat 106

106. Aa hartwegii 113

107. Acianthera casapensis 115

108. Acianthera prognatha 115

109. Acineta cryptodonta 116

110. Acronia archidiaconi 117

111. Acronia bivalvis 118

112. Acronia calamifolia 118

113. Acronia coriacardia 119

114. Acronia phyllocardioides 119

115. Acronia ruberrima 119

116. Acronia semiscabra 120

117. Acronia siphoglossa 120

118. Acronia sp. A 121

119. Anathallis acuminata 122

120. Anathallis sclerophylla 122

121. Brachionidium tuberculatum 123

122. Brassia sulphurea 124

123. Camaridium vestitum 125

124. Cleistes rosea 126

125. Comparettia falcata var. falcata 128

126. Comparettia ottonis 128

127. Corymborkis flava 129

128. Cranichis antioquiensis 130

129. Cranichis diphylla 130

130. Crocodeilanthe elegans 132

131. Crocodeilanthe galeata 133

132. Crocodeilanthe gelida 133

133. Crocodeilanthe moritzii 134

134. Crocodeilanthe stergiosii 134

135. Crocodeilanthe sp. A 134

136. Cyrtochilum cimiciferum 136

137. Cyrtochilum detortum 137

138. Cyrtochilum distans 137

139. Cyrtochilum falcipetalum 137

140. Cyrtochilum megalophium 138

141. Cyrtochilum orgyale 138

142. Cyrtochilum ramosissimum 139

143. Cyrtochilum zebrinum 139
x 

• SMITHSONIAN CONTRIBUTIONS TO BOTANY

144. Dichaea camaridioides 140

145. Dichaea hystricina 141

146. Dichaea latifolia 141

147. Dichaea robusta 142

148. Elleanthus aurantiacus 143

149. Elleanthus columnaris 144

150. Elleanthus confusus 144

151. Elleanthus flavescens 144

152. Elleanthus furfuraceus 145

153. Elleanthus gracilis 145

154. Elleanthus graminifolius 146

155. Elleanthus lupulinus 146

156. Elleanthus wageneri 146

157. Epidendrum alpicola 149

158. Epidendrum andinum 149

159. Epidendrum attenuatum 150

160. Epidendrum caesaris 150

161. Epidendrum cereiflorum 151

162. Epidendrum chioneoides 151

163. Epidendrum curtisii 151

164. Epidendrum dendrobii 152

165. Epidendrum frutex 152

166. Epidendrum guaramacalense 153

167. Epidendrum ibaguense 153

168. Epidendrum jajense 153

169. Epidendrum klotzscheanum 154

170. Epidendrum lacustre 154

171. Epidendrum leucochilum 155

172. Epidendrum pseudocernuum 155

173. Epidendrum repens 156

174. Epidendrum secundum 156

175. Epidendrum unguiculatum 157

176. Epidendrum urichianum 157

177. Epidendrum sp. A 157

178. Epidendrum sp. B 158

179. Epidendrum sp. C 158

180. Fernandezia crystallina 159

181. Fernandezia schultesii 160

182. Gomphichis adnata 161

183. Gomphichis altissima 161

184. Habenaria gollmeri 162

185. Habenaria monorrhiza 163

186. Jacquiniella teretifolia 163

187. Lepanthes glochidea 165

188. Lepanthes scolex 165

189. Lepanthes vareschii 166

190. Lepanthes wageneri 166

191. Lepanthes sp. A 166

192. Lepanthes sp. B 167

193. Lepanthopsis apoda 167

194. Lindleyalis glossopogon 168

195. Lockhartia chocoensis 169

196. Malaxis licatae 170

197. Malaxis nidiae 170
NUMBER 100 • x i

198. Masdevallia gerlachii 172

199. Masdevallia schlimii 172

200. Masdevallia tubulosa 173

201. Maxillaria macrura 174

202. Maxillaria meridensis 174

203. Maxillaria nigrescens 175

204. Maxillaria triloris 175

205. Microchilus paleaceus 176

206. Myoxanthus reymondii 177

207. Odontoglossum dormanianum 178

208. Odontoglossum schillerianum 179

209. Oncidium abortivum 180

210. Ornithidium miniatum 181

211. Ornithidium ruberrimum 181

212. Otoglossum coronarium 182

213. Platystele pisifera 183

214. Pleurothallis dorrii 184

215. Pleurothallis xanthochlora 184

216. Pleurothallopsis striata 185

217. Pleurothallopsis tubulosa 186

218. Ponthieva fertilis 187

219. Prescottia stachyodes 188

220. Prosthechea brachychila 189

221. Prosthechea lindenii 189

222. Psilochilus macrophyllus 190

223. Pterichis multiflora 191

224. Scaphyglottis summersii 192

225. Schlimmia alpina 193

226. Sobralia violacea 194

227. Specklinia endotrachys 195

228. Specklinia zephyrina 195

229. Stelis argentata 197

230. Stelis ascendens 197

231. Stelis atra 198

232. Stelis atroviolacea 198

233. Stelis biserrula 199

234. Stelis chamaestelis 199

235. Stelis humilis 200

236. Stelis hylophila 200

237. Stelis lindenii 200

238. Stelis nitens 201

239. Stelis oblonga 201

240. Stelis pusilla 202

241. Stelis striolata 202

242. Stelis vulcani 202

243. Telipogon andicola 203

244. Telipogon astroglossus 204

245. Telipogon latifolius 204

246. Trichocentrum pulchrum 205

247. Trichopilia laxa 206

248. Trichosalpinx deceptrix 207

249. Trichosalpinx dependens 207

250. Trichosalpinx dunstervillei 208

251. Trichosalpinx pusilla 208
xii • SMITHSONIAN CONTRIBUTIONS TO BOTANY

252. Warreella cyanea 209

253. Agrostis meridensis 212

254. Agrostis mertensii 212

255. Agrostis perennans 212

256. Agrostis pittieri 213

257. Agrostis scabra 213

258. Agrostis venezuelana 214

259. Andropogon bicornis 214

260. Arthraxon hispidus var. hispidus 215

261. Arthrostylidium pubescens 216

262. Arthrostylidium venezuelae 217

263. Aulonemia trianae 218

264. Aulonemia ximenae 218

265. Axonopus compressus 219

266. Bromus catharticus var. catharticus 219

267. Calamagrostis bogotensis 220

268. Calamagrostis planifolia 221

269. Calamagrostis sp. A 221

270. Cenchrus bambusiformis 222

271. Cenchrus clandestinus 223

272. Cenchrus peruvianus 223

273. Chusquea angustifolia 225

274. Chusquea fendleri 225

275. Chusquea mollis 226

276. Chusquea multiramea 226

277. Chusquea serpens 227

278. Chusquea spectabilis 227

279. Chusquea spencei 228

280. Chusquea tessellata 228

281. Cinna poiformis 229

282. Cortaderia hapalotricha 230

283. Danthonia secundiflora subsp. secundiflora 231

284. Dichanthelium acuminatum var. acuminatum 232

285. Didymogonyx geminatum 233

286. Eragrostis tenuifolia 233

287. Eriochrysis cayennensis 234

288. Festuca guaramacalana 235

289. Homolepis glutinosa 236

290. Ichnanthus nemorosus 237

291. Ichnanthus pallens 237

292. Ichnanthus tenuis 237

293. Isachne rigens 238

294. Lasiacis ligulata 239

295. Lasiacis nigra 239

296. Lasiacis procerrima 240

297. Melinis minutiflora 241

298. Ocellochloa pulchella 241

299. Paspalum densum 243

300. Paspalum distichum 243

301. Paspalum inconstans 243

302. Paspalum macrophyllum 244

303. Paspalum paniculatum 244

304. Paspalum penicillatum 244

305. Poa annua 245
NUMBER 100 • xiii

306. Polypogon elongatus 246

307. Rugoloa polygonata 247

308. Setaria parviflora var. parviflora 248

309. Sporobolus indicus 248

310. Smilax domingensis 251

311. Smilax spinosa 251

312. Harperocallis robustior 252

313. Eccremis coarctata 255

314. Xyris subulata var. acutifolia 256

315. Hedychium coronarium 258

316. Renealmia thyrsoidea subsp. thyrsoidea 260

INTRODUCTION

This is the first volume of several that we project will treat all of the ~1,300 species 

of vascular plants that are found in Guaramacal National Park (officially El Parque Nacional “General Cruz Carillo en Guaramacal”). The park was created in 1988 in order to 

protect the Ramal de Guaramacal, an outlier of the more extensive Cordillera de Mérida 

in the Venezuelan Andes and an important watershed for the Boconó and Tucupido reservoirs at the base of the mountains. The park covers 215 km2

 in Trujillo and Portuguesa 

states and is primarily composed of very steep slopes between 1,600 and 3,100 m elevation. The two principal vegetation types found in the park, forest and páramo, have been 

intensively studied from an ecological perspective (Cuello A., 2002, 2010; Cuello A. and 

Cleef, 2009a, 2009b, 2009c, 2011).

Family circumscriptions follow the classification proposed by the Angiosperm Phylogeny Group (2009) as updated by Stevens (2013). In several of the larger families (i.e., 

Araceae, Arecaceae, Bromeliaceae, Orchidaceae, and Poaceae), subfamilial placement of 

a genus is indicated in square brackets following the generic description. This is done to 

suggest at a very crude level the phylogenetic relationships of the genera found within the 

boundaries of the park.

For each species or infraspecific taxon recognized, the accepted name is given in 

bold type. In addition, the basionym (if different) is cited, along with homotypic and 

heterotypic synonyms that have mostly been used in Venezuelan botanical literature. 

These synonymies are intended to help the user understand the application of names 

within Venezuela. Hence, synonymies are not complete, nor are they intended to be complete. References to standard Venezuelan floras such as the Flora de Venezuela (Foldats, 

1969, 1970a, 1970b, 1970c, 1970d; Smith, 1971; Maas, 1982), Flora of the Venezuelan 

Guayana (Berry et al., 1995, 1997, 1998, 1999, 2001, 2003, 2004, 2005), Libro Rojo de 

la Flora Venezolana (Llamozas et al., 2003), and Botánica y Ecología de las Monocotiledóneas de los Páramos en Venezuela (Morillo et al., 2010, 2011) are included as these 

often provide more extensive descriptions and synonymies, as well as illustrations of the 

taxa treated here. Finally, names used in the two earlier enumerations of plants occurring 

in Guaramacal (Ortega et al., 1987; Dorr et al., 2000) are cited, but only when they differ 

from the name recognized in this volume.

Synonymies of orchids are more involved than those of other plant families because 

the Venezuelan orchid literature is rich. Thus, in addition to citing treatments in the Flora 

de Venezuela (Foldats, 1969, 1970a, 1970b, 1970c, 1970d), we also cite descriptions 

and illustrations in Venezuelan Orchids Illustrated (Dunsterville and Garay, 1959, 1961, 

Flora of Guaramacal (Venezuela): 

Monocotyledons

L. J. Dorr

L. J. Dorr, Department of Botany, National Museum of Natural History, MRC-166, Smithsonian 

Institution, P.O. Box 37012, Washington, D.C. 

20013–7012, USA; dorrl@si.edu. Manuscript 

received 8 August 2013; accepted 19 February 

2014.
2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

1965, 1966, 1972, 1976; Dunsterville, 1986), Orchids of Venezuela (Dunsterville and Garay, 1979a, 1979b, 1979c) or its second 

edition (Romero-González and Carnevali Fernández-Concha, 

2000), and Orquídeas Nativas del Táchira (Fernández, 2003).

Distributional statements are intended to be global, but detail by country-level political unit is reserved generally for South 

America. Country-level political units in South America are 

listed from north to south. Distribution within Venezuela is by 

state and federal district (Distrito Federal) and is often grouped 

into the major Andean mountain chains within the country 

(i.e., Cordillera de Mérida, Sierra de Perijá, and Cordillera de la 

Costa). We appreciate that administratively, the Distrito Federal 

was abolished in 1999, but most records of Venezuelan plant 

distribution are based on this former federal district and not 

the Distrito Capital and Vargas state that replaced it. The maps 

are intended to quickly convey distributions within Venezuela, 

and a single dot is placed in the center of the state or federal 

district for which we have a record of occurrence. Localities 

within the park have been simplified from those given on herbarium labels; the north slope faces Boconó and the south slope 

faces the Llanos. The Páramo de Guaramacal is the highest area 

(3,000–3,100 m elevation) within the park and is more or less 

located in the park’s geographical center. The Fila de Agua Fría 

(2,600–2,800 m) is the highest peak to the northeast of the 

Páramo de Guaramacal, whereas the Páramo del Pumar (2,600 

m) is an unusually low and interesting páramo on an exposed 

ridge to the southwest of the Páramo de Guaramacal. Other 

named localities are described in Dorr et al. (2000). Elevations 

cited are for park records and are not to be interpreted as the 

total elevational range of a species or infraspecific taxon. Also, 

elevations cited are mostly taken from labels on specimens and 

rounded to 50 m intervals.

Common names cited in this flora are those that were communicated to us by the people living or working in the park. 

Likewise, the uses of plants that are presented were all provided 

to us by Ramón Caracas, one of the park guards who lives within 

the boundaries of the park.

References are selective. Monographs and revisions are cited 

as are papers that provide the justification for a particular phylogenetic circumscription of a family or genus. Regional treatments 

of families and genera are only cited when they concern taxa 

found in Venezuela or neighboring Colombia.

Abbreviations and Commonly Used Symbols

auct. auctorum (i.e., in the sense of a subsequent author and not the original author)

comb. nov. combinatio nova (i.e., new combination 

of name and epithet)

emend. emendatus (i.e., emended)

f. forma (i.e., form)

fol. folio

ibid. ibidem (i.e., in the same place)

ICN International Code of Nomenclature for 

Algae, Fungi, and Plants (McNeill et al., 

2012)

in litt. in litteris (i.e., in correspondence)

in sched. in schedula (i.e., on a label)

mya million years ago

nom. cons. nomen conservandum (i.e., name conserved by the ICN)

nom. cons. prop. nomen conservandum propositus (i.e., 

proposed conserved name)

nom. illeg. nomen illegitimum (i.e., illegitimate name)

nom. nov. nomen novum (i.e., replacement name)

nom. nud. nomen nudum (i.e., designation of a new 

taxon published without a description or 

diagnosis or reference to a description or 

diagnosis)

nom. & orth. cons. nomen & orthographia conservanda (i.e., 

name and orthography conserved by the 

ICN)

nom. rej. nomen rejiciendum (i.e., name rejected by 

the ICN)

nom. rej. prop. nomen rejiciendum propositus (i.e., proposed rejected name)

pers. comm. personal communication

pers. obs. personal observation

qda. quebrada (i.e., ravine)

qu. quarto

sect. section

s.l. sensu lato (i.e., in the broad sense)

sp., spp. species (singular and plural)

s. str. sensu stricto (i.e., in the strict sense)

subgen. subgenus

subsp. subspecies

t., tt. tabula, tabulae (i.e., plate, plates)

var. variety

vs. versus

♦ signifies that a taxon (or its basionym) 

was described from material collected in 

Guaramacal National Park

• signifies that a taxon is an introduced element in the flora of Guaramacal National 

Park

= heterotypic synonym (i.e., synonym based 

on a different type)

≡ homotypic synonym (i.e., synonym based 

on the same type)

± more or less

× hybrid or times (when used with numbers)

Contributors

Germán Carnevali Fernández-Concha

Centro de Investigacion Cientifica de Yucatán A.C.
NUMBER 100 • 3

Calle 43, No. 130, Col. Chuburná de Hidalgo

97200 Mérida, Yucatán, Mexico

L. J. Dorr

Department of Botany, National Museum of Natural History

Smithsonian Institution, MRC-166

P.O. Box 37012, Washington, D.C. 20013-7012, USA

S. Miguel Niño

Herbario Universitario, BioCentro-UNELLEZ

Mesa de Cavacas 3323, Portuguesa, Venezuela

Anton A. Reznicek

University Herbarium, University of Michigan 

3600 Varsity Dr., Ann Arbor, MI 48109, USA

Basil Stergios D.

Herbario Universitario, BioCentro-UNELLEZ

Mesa de Cavacas 3323, Portuguesa, Venezuela

Mark T. Strong

Department of Botany, National Museum of Natural History

Smithsonian Institution, MRC-166

P.O. Box 37012, Washington, D.C. 20013-7012, USA

SYSTEMATIC TREATMENT

KEY TO THE FAMILIES OF MONOCOTYLEDONS

1a. Plants achlorophyllous; mycoheterotrophic herbs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Burmanniaceae

1b. Plants chlorophyllous; autotrophic herbs, shrubs or trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Plants woody, shrubby or arborescent; stems unbranched; mature leaf blades usually pinnately compound. . . . . Arecaceae

2b. Plants herbaceous or woody, never arborescent; stems branched or not; mature leaf blades simple or, if compound, 

palmately compound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Plants with a Musa- or Zingiber-like habit (i.e., with long, usually oblong, broad leaf blades with a pronounced midrib) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Functional stamens 6 or 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heliconiaceae

 4b. Functional stamens 1 or 1⁄2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 5a. Anthers 2-thecate; flowers zygomorphic; sepals connate below . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 6a. Foliage aromatic; leaves distichous; leaf sheaths usually open; labellum formed by the fusion of 2 

staminodes from the inner floral whorl . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zingiberaceae

 6b. Foliage not aromatic; leaves spirally arranged; leaf sheaths initially closed; labellum formed by the 

fusion of 5 staminodes from both staminal whorls . . . . . . . . . . . . . . . . . . . . . . . . . . . . Costaceae

 5b. Anthers with 1 functional theca, the other anther half petaloid; flowers asymmetric; sepals distinct

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 7a. Flowers borne in mirror-image pairs; stems without mucilage canals; petioles distinctly pulvinate; 

ovary 3-locular or 1-locular by abortion; ovule 1 per locule; seeds arillate (our species) or not . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Marantaceae

 7b. Flowers not paired; stems with mucilage canals; petioles epulvinate; ovary 3-locular; ovules few to 

numerous; seeds not arillate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cannaceae

 3b. Plants without a Musa- or Zingiber-like habit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

 8a. Plants with spathes (i.e., bract-like modified leaves) and flowers aggregated in spadices (i.e., spikes with 

small flowers crowded on a thickened axis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

 9a. Leaf blades plicate and/or bifid apically; spadices subtended by 2–6 mostly caducous spathes (basal 

spathes sometimes persistent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyclanthaceae

 9b. Leaf blades not plicate or bifid apically; spadices subtended by a single mostly persistent spathe . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Araceae

 8b. Plants without spathes and spadices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

 10a. Leaf blades broad, secondary venation ± reticulate; leaf sheaths not obvious; petioles usually well 

defined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

 11a. Petioles forming a sheath that usually terminates in a pair of tendrils; inflorescences umbellate, 

solitary or aggregated in a raceme; fruit a berry . . . . . . . . . . . . . . . . . . . . . . . . . . . Smilacaceae

 11b. Petioles not forming a sheath that terminates in a pair of tendrils; inflorescences spikes or racemes; 

fruit a capsule (our species) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dioscoreaceae
4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

 10b. Leaf blades narrow or broad, secondary venation parallel; leaf sheaths present; petioles not well defined, usually absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

 12a. Flowers greatly reduced; perianth inconspicuous; ovary superior . . . . . . . . . . . . . . . . . . . . . 13

 13a. Plants cushion-like (our species); inflorescences composed of 1 to several terminal, involucrate heads; peduncles enclosed at base by a tubular sheath; flowers usually unisexual 

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriocaulaceae

 13b. Plants not cushion-like; inflorescences and peduncles not as above; flowers unisexual or 

bisexual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

 14a. Perianth biseriate (tepals 6 [4] in 2 whorls), small but evident; flowers not borne in spikes 

or spikelets; ovary 1- or 3-locular; ovules 3 to many; fruit dehiscent . . . . . . Juncaceae

 14b. Perianth not as above; flowers sessile, borne in the axis of scalelike bracts (spikes or 

spikelets); 1-locular; ovule 1; fruit indehiscent . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

 15a. Stems usually triangular in cross section, solid, rarely cylindrical and hollow 

(Eleocharis); leaf sheaths usually closed; flowers subtended by a single scale, usually spirally or less frequently distichously arranged; fruit an achene . . . . . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperaceae

 15b. Stems usually cylindrical in cross section, never triangular, hollow or occasionally solid; leaf sheaths usually open; flowers subtended by a pair of scales, distichously arranged; fruit usually a caryopsis, rarely utricle-like . . . . . . Poaceae

 12b. Flowers generally well developed; perianth (at least some parts) usually petaloid; ovary superior 

or inferior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

16a. Ovary superior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17a. Segments of the perianth differentiated into petals and sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

 18a. Leaves all basal or rosulate, not succulent; leaf sheaths open . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

 19a. Stamens 3; staminodes 3; leaves equitant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xyridaceae

 19b. Stamens 6, in 2 series; staminodes absent; leaves polystichous, often in rosettes or tank-forming (i.e., 

collecting water) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bromeliaceae

 18b. Leaves basal or cauline, not forming rosettes, usually scattered along the stem, ± succulent; leaf sheaths closed, 

sometimes perforated by the peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Commelinaceae

17b. Segments of the perianth similar, not differentiated into sepals and petals . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

 20a. Inflorescences lax, corymbiform panicles; flowers pendulous; tepals blue (rarely white); filaments with a central swelling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthorrhoeaceae

 20b. Inflorescences racemes; flowers ± erect; tepals greenish-yellow; filaments without a central swelling . . . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tofieldiaceae

16b. Ovary inferior or at least partially inferior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

21a. Stamens 1 or 2, adnate to the style in a column; pollen aggregated into pollinia or distinct sticky masses; fruit 

1(3)-locular; seeds minute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orchidaceae

21b. Stamens 3 or 6; not adnate to the style; pollen free (shed as single grains); fruit usually 3-locular; seeds various, not 

minute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

 22a. Stamens 3; leaves equitant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Iridaceae

 22b. Stamens 6; leaves not equitant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

 23a. Leaves cauline or basal and cauline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

 24a. Twining herbs (vine-like); leaves cauline only, resupinate; fruit a semifleshy, leathery capsule; seeds 

usually with a red-orange sarcotesta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alstroemeriaceae

 24b. Terrestrial herbs, not twining or vine-like; leaves basal and cauline, not resupinate; fruit a capsule; 

seeds bicaudate or winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bromeliaceae

 23b. Leaves basal only, often in a rosette . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

 25a. Terrestrial or epiphytic herbs; leaf blades with peltate scales (at least when young) Bromeliaceae

 25b. Terrestrial herbs; leaf blades without peltate scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

 26a. Leaf blades ovate, elliptic, ovate-elliptic or elliptic-lanceolate; flowers white, subpendulous; 

staminal cup conspicuous, 1–1.5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . Amaryllidaceae

 26b. Leaf blades linear to lanceolate; flowers yellow or greenish-yellow, erect or ascending; staminal cup absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypoxidaceae
NUMBER 100 • 5

Alstroemeriaceae

L. J. DORR AND B. STERGIOS

Perennial herbs; twining or erect; mostly glabrous; rhizomatous. Roots with tuber-like swellings for water and carbohydrate storage. Leaves alternate, spirally arranged or distichous, 

often clustered near the ends of elongated stems, simple, entire, 

ovate-lanceolate to lanceolate or linear-lanceolate, membranous, 

papery, subcoriaceous or rarely somewhat fleshy; sometimes appearing petiolate, not sheathing; veins parallel, arcuate-parallel 

or ± reticulate; leaf blades generally semiresupinate or resupinate. Inflorescences terminal (our species) or axillary, helicoid 

cymes in an umbellate arrangement, the flowers often numerous, 

rarely solitary. Involucral bracts foliaceous or reduced. Flowers 

actinomorphic or zygomorphic, epigynous, bisexual. Tepals 3 + 

3 (rarely 4 + 4), petaloid, free, outer whorl often different than 

inner whorl; perigonal nectaries present. Stamens 6 in 2 whorls, 

basifixed or pseudobasifixed; filaments filiform, elongate; anthers 

2-thecate. Ovary syncarpous, inferior, 1⁄2-inferior or superior, 

3-locular, placentation axillary (rarely 1-locular with parietal 

placentation); septal nectaries absent. Style filiform, triquetrous 

at the base, with 3 stigmatic branches at the apex or capitate. 

Tepals, stamens, and style usually caducous. Ovules numerous, 

in 2 rows in each locule, anatropous or hemianatropous. Fruit a 

dry, leathery or somewhat fleshy loculicidal capsule (our species) 

with a circular scar from the tepals or a berry. Seeds globose or 

ovoid, with or without a sarcotesta.

A family of 4 genera and ~200 species that occur in Mexico, 

Central America, the West Indies, South America (all countrylevel political units), New Zealand, and Australia (including Tasmania). Two genera and ~12 species are found in Venezuela.

References. Aagesen and Sanso (2003); Bayer 

(1998); Chacón et al. (2012); Conran and Clifford (1998); Hofreiter (2007); Sanso and Hunziker (1998); Sanso and Xifreda 

(2001).

Our concept of Alstroemeriaceae includes Luzuriaga Ruiz 

& Pav. and Drymophila R. Br., which have been segregated as 

Luzuriagaceae. Alstroemeriaceae is principally American with a 

few exceptions; Bomarea Mirb. and Alstroemeria L. are wholly 

American, three species of Luzuriaga are found in Chile and one 

in New Zealand, and one species of Drymophila is found in Australia and another one in Tasmania.

Bomarea Mirb.

Bomarea Mirb., Hist. Nat. Pl. 9: 71. 1804.

Twining herbs; prostrate, climbing, scandent or erect; stems 

often tortuous; roots fibrous, often tuberous at the tip. Leaves resupinate; petioles relatively short; leaf blades ovate-lanceolate to 

linear-lanceolate, margins entire, membranous, papery or subcoriaceous, nerves parallel, glabrous, hirsute or glandular-puberulent. 

Inflorescences umbellate, simple or compound, few- to manyflowered. Involucral bracts foliaceous or greatly reduced. Flowers actinomorphic or slightly zygomorphic; often nutant. Perianth 

funnelform or tubular, whitish to orange, pink, red, purple or 

greenish; outer tepals oblong, uniform, concolorous, sometimes 

shorter than the internal ones, hooded or not; inner tepals clawed 

or spathulate, lowermost tepal smaller than the other tepals, differently colored, often maculate. Stamens erect or curved, as long 

as the perianth; filaments glandular-puberulent or glabrous. Ovary 

inferior (our species), 3-locular. Fruit a semifleshy, leathery capsule. Seeds numerous, usually with a red-orange sarcotesta.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, the Guianas, Ecuador, 

Peru, Bolivia, Brazil, Chile, Paraguay, and Argentina); introduced 

into New Zealand where 1 species is now an invasive weed. The 

genus includes 100–110 species, of which ~10 occur in Venezuela. These Venezuelan species are almost exclusively montane; 

only Bomarea edulis (Tussac) Herb. and B. ovata (Cav.) Mirb. 

are found at lower elevations.

References. Alzate et al. (2008a, 2008b); Baker 

(1888); Garbiso and Estrada (2001); Hofreiter (2006); Hofreiter 

and Tillich (2002).

Bomarea is closely related to Alstroemeria but can be separated by its actinomorphic (sometimes slightly zygomorphic) 

flowers, straight filaments, leathery to somewhat fleshy capsules 

or berrylike fruits, and sarcotesta. Alstroemeria has zygomorphic flowers, curved filaments, dry capsules with explosive dehiscence, and a dry seed coat.

Baker (1888) proposed a classification of Bomarea based 

on morphology in which he recognized three subgenera. Hofreiter and Tillich (2002) added a fourth subgenus, also based 

on morphology. However, Alzate et al. (2008a) recovered three 

clades in a molecular phylogenetic analysis of the genus, none 

of which correspond to the subgeneric groupings based on 

morphology.

KEY TO THE SPECIES OF BOMAREA

1a. Inflorescences compound umbels, rays 1–3-branched; bracteoles (floral bracts) present; tepals <3 cm long . . . . . . . . . 2

2a. Cauline leaves 0.8–1.6(–2.5) cm wide, glabrous above, glaucous below with a whitish, scalelike pubescence; rays 

1-branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. amilcariana

2b. Cauline leaves 2–2.5(–3) cm wide, glabrous above and below or with hairs on the nerves below; rays 1–3-branched

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. edulis

1b. Inflorescences simple umbels, rays unbranched; bracteoles (floral bracts) absent; tepals >4 cm long . . . . . . B. salicifolia
6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

♦ Bomarea amilcariana Stergios & Dorr, Acta Bot. Venez. 26: 

32, figs. 1, 2. 2003 [2004]; Briceño, in Morillo et al., Bot. 

Ecol. Monocot. Páramos Venez. 1: 51–52, fig. 1. 2011.

FIGURES 1, 23A

Bomarea sp. B; Dorr et al., Contr. U.S. Natl. Herb. 40: 36. 2000 [2001].

Bomarea pauciflora auct., non (Kunth) Herb.; Garbiso and Estrada, PlantULA 3: 25–27. 2001, pro parte, excluding type.

Twining herbs; prostrate, climbing or scandent. Leaf blades 

linear-lanceolate (rarely narrowly ovate-lanceolate), 9–11(–15) × 

0.8–1.6(–2.5) cm, subcoriaceous, glabrous above, glaucous below 

FIGURE 1. Alstroemeriaceae, Bomarea amilcariana Stergios & Dorr. A. Habit. B. Leaf undersurface showing scale-like pubescence. C. Detail 

of multicellular pubescence on umbel rays. D. Perianth. E. Outer tepal (abaxial view). F. Inner tepal (abaxial view). G. Fruit. (A–F, Stergios & 

Dorr 8387; G, Dorr & Barnett 8073.)
NUMBER 100 • 7

with a whitish, scalelike vestiture; petioles to 7 mm long. Involucral bracts foliaceous, 2–5 cm long. Umbels lax, ~10–11 cm in diameter; rays 6–12(–14), 1-branched, (3–)5–8.5 cm long, ± densely 

hirsute-pubescent; floral bracteoles 5–7 mm long. Tepals equal or 

subequal; outer tepals obovate to broadly spathulate, 2–2.3 × 1 

cm, dark or bright red with yellow on the midvein distally; inner 

tepals spathulate, 2.3–2.6 × 0.7–0.8 cm, green with brown maculations distally. Capsules triangular-turbinate, 1–2 cm in diameter.

Endemic to the Andes of Venezuela (Trujillo; Map 1). Known 

from cloud forest and páramo on both slopes of Guaramacal as 

well as from the Páramo del Pumar and the Fila de Agua Fría; 

(1,600–)1,900–3,100 m.

Bomarea edulis (Tussac) Herb., Amaryllidaceae 111. 1837; 

Meerow, in Berry et al., Fl. Venez. Guayana 6: 5, fig. 2. 2001. 

Alstroemeria edulis Tussac, Fl. Antill. 109, t. 14. 1808.

♦Bomarea truxillensis Stergios & Dorr, Acta Bot. Venez. 26: 36. 2003 [2004].

Bomarea obovata auct., non Herb.; Dorr et al., Contr. U.S. Natl. Herb. 40: 

36. 2000 [2001].

Twining herbs; scandent, prostrate or creeping. Leaf blades 

lanceolate to linear-lanceolate, (5–)11–12(–20) × 2–2.5(–5) cm, 

membranous or papery, glabrous above and below or with hairs 

on the nerves below; petioles 5–8 mm long. Involucral bracts 

foliaceous, 2–4 × 0.3–0.5 cm. Umbels lax, ~9–11 cm in diameter; 

rays 3–23, 1–3-branched, 3–15(–26) cm long; secondary and tertiary rays always shorter than primary ones; floral bracteoles to 

1.5 cm long. Tepals subequal to unequal; outer tepals broadly 

ovate, 1.9–3 × 0.6–1.3 cm, pink, pink-orange or pink-purple; 

inner tepals spathulate, 2–2.7(–3) × 0.5–0.7(–1) cm, white or 

green with a subapical row of dark maculations. Capsules turbinate or 3-angled, 2–3 cm in diameter.

Found in Mexico, Central America, the Greater Antilles, 

and South America (Colombia, Venezuela, the Guianas, Peru, 

Bolivia, Brazil, Paraguay, and Argentina). In Venezuela, found in 

Amazonas, Aragua, Barinas, Bolívar, Cojedes, Distrito Federal, 

Falcón, Lara, Mérida, Miranda, Portuguesa, Sucre, Táchira, and 

Trujillo (Map 2). In the park, found on both slopes of Guaramacal; 1,900–2,900 m.

This species has the widest distribution in the genus and is 

morphologically variable, two factors that have contributed to 

an extensive synonymy. The tuberous roots have been used as 

food, and this may be the reason for its extensive geographic 

range.

Several species are confused with Bomarea edulis. Nearly 

closed, cucullate flowers distinguish B. obovata Herb. from 

B. edulis, which has funnel-shaped flowers. Also, in herbaria, 

B. edulis often is confused with B. ovata, but the outer tepals 

are broadly ovate in the former and narrowly ovate in the latter. Also, the ovary is glabrous or nearly glabrous in B. edulis

but densely pubescent in B. ovata. Bomarea ovata appears to 

be confined to Peru, Bolivia, and Argentina, whereas B. edulis is 

more widespread.

Bomarea salicifolia Killip, Natl. Hort. Mag. 15: 120. 1936.

Twining herbs; scandent or prostrate. Leaf blades linearlanceolate, 11–13(–24) × 0.8–0.9(–1.5) cm, subcoriaceous, glabrous above and below; petioles 8–10 mm long. Involucral bracts 

± dimorphic, outer bracts subfalcate, 4–5 cm long, inner ones 

linear-lanceolate, 1–1.5 cm long. Umbels congested, ~20 cm in 

diameter, with 10–40 rays, rays 5–5.5 cm long, unbranched; floral bracteoles absent. Tepals subequal; outer tepals oblanceolate, 

4.8–5.1 × 0.8–1.2 cm, pink; inner tepals spathulate, ~4.5 × 1 cm, 

MAP 1. Bomarea amilcariana occurrence in Venezuela. MAP 2. Bomarea edulis occurrence in Venezuela.
8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

green with purple spots near the margins and distally. Capsules 

not seen.

Endemic to the Andes of Venezuela (Trujillo; Map 3). In the 

park, found along the Boconó–Guaramacal road; 2,600 m.

This species is rarely collected. It was first gathered in 1846 

at an unspecified locality in Trujillo state, described 90 years 

later, and only collected a second time (our voucher, Niño et al. 

1390) in 2000. When Killip (1936) described Bomarea salicifolia, he misinterpreted its habit as erect rather than prostrate, 

which is understandable given the limited amount of material 

that was available to him.

• Amaryllidaceae

L. J. DORR AND B. STERGIOS

Perennial, mostly geophytic herbs; terrestrial (our species), 

sometimes aquatic or epiphytic; bulbs tunicate. Leaves annual 

or persistent, distichous or spirally arranged; leaf blades sessile 

and linear or lorate, or petiolate and lanceolate to widely elliptic; 

sometimes with a sheath at the base and forming an aerial pseudostem; usually glabrous. Inflorescences scapose, pseudoumbellate (i.e., reduced helicoid cymes); scapes terminated by 2 or more 

spathe-like, usually marcescent bracts (rarely bracts absent); inner 

bracteoles usually present and successively shorter and narrower. 

Flowers 1 to many, usually showy, sessile or pedicellate, actinomorphic or zygomorphic, protandrous, each flower subtended by 

a bracteole, bisexual. Perianth crateriform, salverform, funnelform, tubular or ventricose, composed of 6 tepals in 2 whorls (3 + 

3), connate below to form a tube or rarely free to the base; inner 

tepals shorter than outer ones; outgrowth of perianth sometimes 

present, forming a conspicuous crown or relatively inconspicuous 

rim of scales at the throat. Stamens 6 (3 + 3; rarely more), subequal, inserted in the throat of the perianth or below; filaments 

variously connate; anthers usually dorsifixed, introrse, dehiscing 

longitudinally or rarely poricidally. Style filiform. Stigma capitate, 

3-lobed or deeply trifid. Ovary syncarpous, inferior, 3-carpellate, 

3-locular (rarely 1-locular); septal nectaries present; placentation 

axile or basal; ovules anatropous. Fruit a capsule, loculicidally dehiscent or sometimes indehiscent, rarely berrylike. Seeds globose 

or subglobose, fleshy or hard, or flattened and winged, usually 

with a black or brown phytomelanous seed coat.

A family of ~73 genera and ~1,600 species found in tropical 

and temperate regions worldwide. Nine genera and 25 native 

and naturalized species are reported to occur in Venezuela.

References. Kubitzki (1998a); Meerow and Snijman (1998); Meerow et al. (1999, 2000); Rahn (1998).

Our concept of Amaryllidaceae includes Alliaceae and Agapanthaceae, but our family description emphasizes characters of 

taxa most likely to be encountered in the neotropics. Amaryllidaceae is not well represented in Venezuela, although there are 

a number of taxa cultivated as ornamentals, some of which have 

escaped and become naturalized.

• Eucharis Planch. & Linden

Eucharis Planch. & Linden, in Linden, Cat. Pl. Exot. 8: 3. 1853, nom. cons.

Perennial, geophytic herbs; bulbs usually offsetting vigorously. Leaves persistent, glabrous; petioles subterete, winged 

distally by attenuation of the base of the leaf blade; leaf blades 

ovate, elliptic, ovate-elliptic or elliptic-lanceolate, margins frequently undulate, membranous or papery, lustrous, parallelveined, smooth or plicate between the veins. Inflorescences 

scapose, pseudoumbellate (i.e., 1 to several reduced helicoid 

cymes); scapes solid, terminating in 2 green or greenish-white 

ovate-lanceolate, marcescent bracts. Flowers 2–12, pedicellate 

or rarely subsessile, pendulous or declinate, white, sometimes 

fragrant; perianth campanulate or crateriform; perianth tube cylindrical, dilating abruptly above the middle; perianth limb of 

6 tepals in 2 series, spreading widely from the throat or partly 

imbricate; outer tepals longer than inner tepals. Stamens 6, variously connate below; free filament linear, subulate or otherwise 

petaloid; anthers oblong to linear, subbasifixed or dorsifixed, 

eventually versatile, introrse. Style filiform, included within, 

equal to or exserted from the staminal cup. Stigma obtusely 

3-lobed. Ovary inferior, green or white, ellipsoid-oblongoid, 

sometimes trigonous, 3-locular; ovules (2–)7–20 per locule; placentation axile. Fruit a 3-lobed loculicidal capsule, thin walled or 

leathery, green or bright orange. Seeds few per locule, globose or 

ellipsoid, lustrous (rarely dull) black, dark brown or blue.

A neotropical genus of 17 species and various natural hybrids found in Central America, the West Indies, and South 

America (Colombia, Venezuela, Trinidad and Tobago, Suriname, 

Ecuador, Peru, Bolivia, and Brazil). In Venezuela, 2 species are 

cultivated or naturalized.

MAP 3. Bomarea salicifolia occurrence in Venezuela.
NUMBER 100 • 9

References. Meerow (1989); Meerow and Deghan 

(1984).

The genus Eucharis as circumscribed by Meerow and Snijman (1998) and Meerow (1989) may not be monophyletic 

(Meerow et al., 2000).

• Eucharis amazonica Linden ex Planch., Fl. Serres Jard. Eur. 12 

[sér. 2, 2]: 69, tt. 1216, 1217. 1857.

Herbs, 40–50 cm tall; bulbs 3–5 cm in diameter, tunics 

brown. Leaves 2–4; petioles slender, 15–20(–50) cm long; leaf 

blades elliptic, 15–17(–50) × 8–9 cm, bases attenuate, margins ± 

undulate, apices acuminate, shallowly or inconspicuously plicate, 

dark green above, lighter green below. Scapes 28–32(–80) cm 

long, narrowing distally. Flowers 4–6(–8), subpendulous, white, 

sweet scented; pedicels 15–17(–25) mm long, slightly curved; perianth tube strongly curved, cylindrical below, dilating in upper 1⁄3, 

white; perianth limb spreading widely from the throat of the tube, 

9–10 cm wide; tepals ovate; outer tepals apiculate, 4–4.5 × 2–3 

cm; inner tepals acute, 3.5–4 × 2.5–3.5 cm. Staminal cup widely 

cylindrical, 1–1.5 cm long; stamens with 2 obtuse lobes, 1 on each 

side of the subulate free filament. Style white, exserted from the 

staminal cup. Ovary oblong-ellipsoid, 0.7–1.3 cm long; ovules 

9–15 per locule, superposed. Fruit and seed unknown.

Native to northern Peru but widely cultivated and now 

naturalized in Central America (Panama), the West Indies, and 

South America. In Venezuela, known from gardens in Aragua, 

Carabobo, Distrito Federal, Nueva Esparta, and Trujillo; naturalized in Trujillo, at least (Map 4). In the park, found near the 

Laguna de Aguas Negras; ~1,850 m.

Eucharis amazonica is widely and incorrectly known in the 

horticultural trade as E. grandiflora Planch. & Linden. Meerow 

(1989) considered this latter taxon, which has not been found 

in Guaramacal, to be a hybrid (i.e., E. ×grandiflora Planch. & 

Linden) that originated in Colombia. The parents of this hybrid 

are E. moorei (Baker) Meerow and E. sanderi Baker.

Araceae

L. J. DORR AND B. STERGIOS

Terrestrial, epiphytic, hemiepiphytic or lithophytic herbs, 

rarely aquatic; latex milky or watery; rhizomes, corms or stolons 

present; stems aerial and erect, climbing or scandent. Leaves spirally arranged or sometimes distichous; cataphylls (i.e., bladeless 

leaves corresponding in appearance to petiole sheaths) usually 

present, deciduous or persistent, intact or weathering into fibers; 

petioles often geniculate apically; blades simple, lobed or compound, sometimes perforate, margins entire, usually glabrous, 

primary lateral nerves pinnate, lesser nerves striate or reticulate. Inflorescences 1 to several terminal (sometimes appearing 

axillary) spadices (i.e., densely congested, unbranched spikes of 

minute, ebracteolate flowers), each spadix subtended or enveloped by a single, often colored spathe (i.e., bract-like modified 

leaf), free or sometimes partially adnate to the spadix, spreading, 

reflexed or convolute, sometimes constricted below the middle 

and differentiated into a proximal tube and distal blade, often 

persistent. Flowers bisexual or unisexual, when unisexual then 

plant usually monoecious and staminate flowers distal to the pistillate ones; staminate and pistillate flowers contiguous or separated by sterile flowers. Perianth absent or present; when present 

composed of 4–6 free or rarely united tepals. Stamens 2–6(–9), 

free or united in synandria (i.e., staminate flowers composed of 

connate stamens). Pistil 1, free. Ovary usually superior, 1–3(to 

many)-loculed; ovules 1 to many; placentation axile, parietal, 

basal or apical. Style 1, inconspicuous. Stigma ± discoid. Fruit a 

berry, 1- to many-seeded. Seeds various; testa thin to thick.

A cosmopolitan family of ~120 genera and ~3,800 species, 

with its greatest diversity in the tropics. The family is found in 

all country-level political units in the Americas except for Greenland. Twenty-four genera and ~290 species occur in Venezuela.

References. Bunting (1979); Cabrera et al. (2008); 

Croat and Lambert (1986); Cusimano et al. (2011); Madison 

(1978b); Mayo et al. (1997, 1998); Nauheimer et al. (2012); Nicolson (1982); Rothwell et al. (2004); Schott (1860); Wilson (1960).

The family circumscription adopted here includes Lemnaceae, which long was considered to be a separate family composed exclusively of aquatic plants (see, e.g., Landolt in Hokche 

et al., 2008). Molecular data, however, indicate that these floating aquatics belong in Araceae (Rothwell et al., 2004), and they 

are now treated as a distinct clade (subfamily Lemnoideae) sister 

to the “true” Araceae (Cusimano et al., 2011).

A subfamily classification based on nonmolecular characters proposed by Mayo et al. (1997) recognized seven subfamilies (not including Lemnoideae), and it was largely confirmed 

in a molecular analysis by Cabrera et al. (2008). However, 

MAP 4. Eucharis amazonica occurrence in Venezuela.
1 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

reanalysis of molecular data by Cusimano et al. (2011) with 

more complete gene sampling identified 44 larger clades, most 

of which are informally named. These clades undoubtedly will 

eventually serve as the basis for a new subfamily classification.

KEY TO THE GENERA OF ARACEAE

1a. Spathe undifferentiated; flowers bisexual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Spathe persistent after anthesis; flowers with 4 tepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anthurium

2b. Spathe marcescent or deciduous soon after anthesis; flowers achlamydeous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Leaf blade entire, never perforate; ovules numerous per locule . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhodospatha

 3b. Leaf blade often perforate, sometimes perforations laterally open to the margins; ovules 2 per locule . . . . . . . . 

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monstera

1b. Spathe differentiated into a convolute tube proximally and an expanded blade distally; flowers unisexual . . . . . . . . . . 4

4a. Climbing, hemiepiphytic or epiphytic herbs; spathe and/or spadix secreting resin at anthesis; staminate flowers with 

2–6 free stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Philodendron

4b. Terrestrial ± arborescent herbs (our species); spathe and spadix dry at anthesis; staminate flowers with 4–6 connate 

stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthosoma

Anthurium Schott

Anthurium Schott, Wiener Z. Kunst 1829: 828. 1829.

Epiphytic, hemiepiphytic, lithophytic or terrestrial herbs; 

rarely rhizomatous; stems erect, creeping or climbing; laticifers absent. Leaves often clustered near the apex of stems; cataphylls entire (membranous) or rotting and forming a fibrous mass, usually 

± persistent; petioles geniculate apically, terete to angled; sheaths 

conspicuous in juvenile leaves and very short in mature ones; 

blades simple, dissected or compound, variously shaped, linear to 

orbicular in outline, primary lateral veins pinnate, forming 1 or 

more submarginal collective veins, basal ribs often present in cordate leaves, higher-order venation reticulate, coriaceous, membranous or stiff and brittle. Inflorescences solitary. Peduncles elongate. 

Spathe simple, usually linear to elliptic or ovate, erect, spreading 

or reflexed, not enclosing the spadix, usually persistent, sometimes 

marcescent or deciduous. Spadix sessile to long-stipitate, cylindrical to conical, less commonly clavate or globose, very short to very 

long (>1 m long). Flowers bisexual, perigoniate (i.e., with a floral 

envelope in which there is no differentiation between sepals and 

petals); tepals 4, fornicate (i.e., with scalelike appendages) in 2 

decussate whorls. Stamens 4, free. Ovary ovoid to oblong or obovoid, 2-locular; ovules 1 or 2(rarely more) per locule, anatropous 

to campylotropous. Stylar region inconspicuous. Stigma small, 

subcapitate. Berry globose to oblong-fusiform. Seeds 2–4, variously colored; testa thin, smooth or subverrucose, often sticky.

A neotropical genus of 900–1,000 species found in Mexico, Central America, the West Indies, and South America (all 

country-level political units except Uruguay); introduced in tropical Asia. Seventy-five species are found in Venezuela.

References. Carlsen (2011); Croat (1985, 1991); 

Croat and Bunting (1979); Croat and Carlsen (2013); Croat and 

Sheffer (1983); Engler (1898, 1905); Madison (1978a); Sheffer 

et al. (1980).

The genus Anthurium is monophyletic (Carlsen, 2011). Anthurium and several smaller genera found in Madagascar, Asia, 

and Australasia are currently placed in a clade that corresponds 

to the subfamily Pothoideae (Cusimano et al., 2011), which is 

sister to the Monsteroideae. In the scheme outlined by Cusimano 

et al. (2011) these two subfamilies form a larger clade that is 

informally named the bisexual climbers clade.

Infrageneric classification of Anthurium is less well resolved. 

Carlsen (2011), using molecular evidence, determined that for 

the most part the sectional classification of Anthurium proposed 

by Croat and Sheffer (1983) is artificial and does not reflect the 

phylogeny of the genus.

KEY TO THE SPECIES OF ANTHURIUM

1a. Leaves palmately compound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. eminens

1b. Leaves simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Bases of leaf blades obtuse to subtruncate or slightly cordate, not lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Leaf blades dark glandular-punctate above and below . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Stems and leaves tightly adhering to substrate (usually tree trunks); leaf blades 1.2–3(–5.5) cm wide; spadix 

<2 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. scandens

 4b. Stems and leaves not tightly adhering to substrate, usually erect; leaf blades 3.5–18 cm wide; spadix >2.5 cm 

long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
NUMBER 100 • 1 1

 5a. Leaf blades ovate to broadly ovate, 11–18 cm wide; petioles much longer than leaf blades . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. bernardii

 5b. Leaf blades lanceolate to narrowly ovate, 4–5 cm wide; petioles shorter than leaf blades . . . . A. gehrigeri

 3b. Leaf blades not dark glandular-punctate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 6a. Leaf blades brownish glandular-punctate above and below (punctations often difficult to see); petioles much 

longer than leaf blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. longegeniculatum

 6b. Leaf blades without glandular punctations, dark or otherwise; petioles ± equal to or shorter than leaf blades 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 7a. Leaf blades <15 cm wide, collective vein continuous from near base to apex; cataphylls deciduous; 

bases of leaf blades cuneate to narrowly attenuate, truncate or rounded; petioles subterete, <0.5 cm in 

diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. amoenum

 7b. Leaf blades >20 cm wide, collective vein not continuous; cataphylls persisting, fibrous; bases of leaf 

blades long-attenuate; petioles sulcate-triangular, ≥1 cm in diameter . . . . . . . . . . . . . A. crassinervium

2b. Bases of leaf blades lobed; lobes unequal, separated by a sinus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

 8a. Spathe lanceolate to narrowly or broadly lanceolate, recurved at anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

 9a. Spathe bright or brick-red (less commonly reddish-pink); peduncles 14–17.5(–36) cm long . . . . A. julianii

 9b. Spathe green; peduncles 32–53 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. smaragdinum

 8b. Spathe ovate or ovate-lanceolate, erect, not recurved at anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

 10a. Cataphylls remaining intact; leaf blades ovate to broadly ovate, never constricted . . . . . . . . . . . . . . . . 11

 11a. Spathe white or pinkish-white; spadix shorter than spathe . . . . . . . . . . . . . . . . . A. nymphaeifolium

 11b. Spathe light greenish-cream to greenish-yellow; spadix longer than spathe . . . . . A. ramoncaracasii

 10b. Cataphylls disintegrating into a mass of fibers (at least below); leaf blades obpyriform to triangular-ovate, 

constricted or not . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

 12a. Leaf blades obpyriform, slightly to strongly constricted; sinus between basal lobes parabolic to 

broadly parabolic, lobes divergent, never overlapping . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. ginesii

 12b. Leaf blades triangular-ovate, not constricted; sinus between basal lobes hippocrepiform to spathulate, lobes not divergent, sometimes overlapping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. nubicola

Anthurium amoenum Kunth & C. D. Bouché, Index Seminum 

[Berlin] 1848: 11. 1848.

FIGURE 23B

Anthurium humile Schott, Prodr. Syst. Aroid. 442. 1860. Anthurium amoenum var. humile (Schott) Engl., Bot. Jahrb. Syst. 25: 380. 1898; Dorr et 

al., Contr. U.S. Natl. Herb. 40: 37. 2000 [2001].

Terrestrial, climbing or epiphytic herbs; stems typically 

creeping. Leaves simple; cataphylls deciduous; petioles 25–

38(–50) cm long, ± equal in length to leaf blades; leaf blades 

elliptic to broadly elliptic or obovate, 28–42 × (6.5–)9–16 cm, 

bases cuneate to narrowly attenuate, truncate or rounded, apices 

long-acuminate, apiculate, primary lateral veins ascending, collective vein slightly sinuate, weakly loop connected at primary 

lateral veins, 3–8 mm from the margin, membranous, without 

glandular punctations. Inflorescences erect, slightly longer than 

petioles. Peduncles 20–50(–68) cm long. Spathe broadly lanceolate, 4–9 × 1–1.7 cm, recurved, green or pale green. Spadix not 

stipitate, cylindrical, slightly tapering to the apex, 4–8.5(–11) cm 

long, green or greenish-yellow.

Restricted to South America (Colombia, Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, collected in the Andes 

(Barinas, Lara, Mérida, Portuguesa, and Trujillo) and the Cordillera de la Costa (Aragua, Distrito Federal, Falcón, Miranda, and 

Yaracuy; Map 5). In the park, found in montane forest on both 

slopes of Guaramacal; (1,600–)1,800–2,100 m.

MAP 5. Anthurium amoenum occurrence in Venezuela.
1 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Engler (1898) recognized two varieties of this species, which 

he distinguished on relatively minor differences in leaf shape. Although we accepted this distinction earlier (Dorr et al., 2000), 

it now appears to us that there is only a single, variable, and 

widespread Andean species; differences in leaf base shapes that 

we cited as being diagnostic occur within populations.

Anthurium bernardii Croat, Aroideana 8: 120. 1985 [1986].

Anthurium bredemeyeri auct., non Schott; Ortega et al., BioLlania 5: 33. 

1987.

Anthurium fernandezii auct., non Croat; Dorr et al., Contr. U.S. Natl. Herb. 

40: 37. 2000 [2001].

Terrestrial, climbing or epiphytic herbs; stems erect, to 20(–

30) cm tall. Leaves simple; cataphylls persisting as a network of 

fibers that surround the lowermost internodes; petioles 28–48 cm 

long, much longer than leaf blades; leaf blades ovate to broadly 

ovate, 22–32 × 11–18 cm, bases acute to rounded, apices acuminate, primary lateral veins pinnate-parallel, collective vein straight 

or weakly loop connected at primary lateral veins, 2–8 mm from 

margin, subcoriaceous, blackish glandular-punctate above and 

below. Inflorescences erect, shorter than petioles. Peduncles 18–35 

cm long. Spathe lanceolate, to 1 × 6 cm, erect to reflexed, clasping 

at base, dark green outside to reddish-green or reddish-brown inside. Spadix sessile to short-stipitate, cylindrical, slightly tapering 

apically, 5.5–10 cm long, reddish-brown to purple.

Endemic to the Cordillera de Mérida in Venezuela (Barinas, 

Mérida, and Trujillo; Map 6). In the park, found in montane 

forest and cloud forest on both slopes of Guaramacal, including the karst area above the Río Amarillo; (1,200–)1,600–2,350

(–2,700) m.

This species is superficially similar to Anthurium smithii

Croat, which has a more extensive range in Andean Venezuela 

and Colombia, but the two species are easily distinguished because the latter has dark glandular punctations on only the lower 

surface of the leaf blades.

Anthurium crassinervium (Jacq.) Schott, Wiener Z. Kunst 1829: 

828. 1829. Pothos crassinervius Jacq., Collectanea 4: 122. 

1790 [1791] (“crassinervia”).

Terrestrial, lithophytic or epiphytic herbs; stems short, 

leaves erect to spreading from a rosette-like base. Leaves simple; 

cataphylls persisting ± intact, weathering into a network of fibers; petioles 3–35 × 0.5–1 cm, generally quadrangular in cross 

section, thicker than broad, sulcate; leaf blades oblanceolate, 25–

150 × 10–50 cm, bases acute to obtuse, margins undulate, apices 

gradually acuminate or rounded, primary lateral veins ascending, collective vein arising in the upper 1⁄3–1⁄4 of blade, coriaceous. 

Inflorescences erect to spreading. Peduncles 20–98 × 0.5–1 cm, 

usually longer than petioles. Spathe lanceolate, 8–15 × 1.5–2 cm, 

spreading to reflexed, green, sometimes suffused with purple, 

deciduous. Spadix sessile to short-stipitate, gradually attenuate, 

(5–)12–35 × 0.7–1.5(–2.5) cm, dark purple or greenish-purple. 

Infructescence pendent; berries red.

Native to Mexico, Central America, and South America (Colombia, Venezuela, Curaçao, Guyana, and Peru); introduced in 

Australia (Queensland). Widespread in Venezuela (Amazonas, Anzoátegui, Aragua, Bolívar, Carabobo, Distrito Federal, Falcón, Lara, 

Mérida, Miranda, Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, and Zulia; Map 7). Found in montane forest and 

cloud forest on both slopes of Guaramacal; 1,850–1,950 m.

This species is part of a complex that has an extensive geographical distribution. If considered in its narrowest sense (e.g., 

Mayo, 1982), the range of our taxon would be restricted to Colombia and western Venezuela.

MAP 6. Anthurium bernardii occurrence in Venezuela. MAP 7. Anthurium crassinervium occurrence in Venezuela.
NUMBER 100 • 1 3

Anthurium crassinervium s.l. is cultivated and available in 

the horticultural trade as bird’s nest anthurium.

Anthurium eminens Schott, Oesterr. Bot. Wochenbl. 5: 273. 

1855; Bunting, in Berry et al., Fl. Venez. Guayana 2: 608. 

1995.

Anthurium wittianum Engl., nom. nud., in sched.

Anthurium ascensus G. S. Bunting, nom. nud., in sched.

All Venezuelan material is referred to the nominate 

subspecies.

Anthurium eminens subsp. eminens.

Epiphytic, hemiepiphytic or scandent herbs; stems to 10 m 

long. Leaves palmately compound; cataphylls disintegrating into 

a network of pale, persistent fibers; petioles 50–75 cm long, canaliculate; leaflets 11–15, radiating from a central nexus; petiolules 

4–12 cm long; leaflet blades narrowly oblong to oblong, 5–10 × 

20–40 cm, somewhat unequal in size with lateral leaflets smallest, bases cuneate to truncate, apices long-acuminate, almost 

caudate, primary lateral veins parallel-ascending, collective vein 

arising near base and extending to apex, 3–4 mm from margin, 

coriaceous. Inflorescences ± ascending. Peduncles 30–35 cm long, 

shorter than petioles. Spathe linear-lanceolate, 20–50 × (0.1–)1–3 

cm, widest near base and tapering to a fine point apically, reflexed 

and twisted, green, sometimes suffused with purple, persistent. 

Spadix short-stipitate, cylindrical, 25–45 cm × 6–12 mm, expanding in fruit, dark dull green or bright purple fading to brownishgray. Berries purple, often found only at base of spadix.

Found in South America (Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, and Brazil). In Venezuela, known 

from the Andes (Barinas, Lara, Portuguesa, and Trujillo), the 

Cordillera de la Costa (Miranda), and the Venezuelan Guayana 

(Amazonas and Bolívar; Map 8). Found in forest on south-facing 

slopes in La Divisoria de la Concepción, El Santuario, and other 

localities in the park; (1,500–)1,800–2,100 m.

Common Name. Guajesito.

Anthurium eminens subsp. longispadix Croat & M. M. 

Mora, described from and evidently endemic to the Chocó region of Colombia, is distinguished by its long, grayish-green spadix that is more slender and about twice as long as that of the 

nominate subspecies.

Carlsen (2011) demonstrated with molecular data that the 

palmately divided leaf morphology in Anthurium evolved independently at least three times. Nonetheless, most of the species with 

palmately divided leaves, including A. eminens, fall into a single 

clade, which is recognized as A. sect. Dactylophyllium (Schott) 

Engler emend. Croat & Carlsen (Croat and Carlsen, 2013).

Anthurium gehrigeri Croat, Aroideana 8: 129. 1985 [1986]; 

Morillo and Briceño, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 1: 69. 2011.

Epiphytic or climbing herbs; stems erect to ascending. 

Leaves simple; cataphylls persisting as a network of fibers covering the stem and internodes; petioles 9–18(–23) cm long, generally shorter than leaf blades; leaf blades lanceolate to narrowly 

ovate, 14–28.5 × 3.5–4.5(–6.5) cm, bases acute, apices acuminate to long-acuminate, primary lateral veins ascending, collective vein straight or weakly arcuate, 5–7 mm from margin, 

subcoriaceous, blackish glandular-punctate above and below. 

Inflorescences erect. Peduncles (8.5–)18–30 cm long, longer than 

petioles (shorter than petioles and blades combined). Spathe lanceolate, 2–5 × 0.5 cm, greenish, recurved. Spadix short-stipitate, 

narrowly cylindrical, (3–)5–7 cm long, greenish.

MAP 8. Anthurium eminens subsp. eminens occurrence in 

Venezuela. MAP 9. Anthurium gehrigeri occurrence in Venezuela.
1 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Endemic to Venezuela, where it is found in the Andes 

(Mérida and Portuguesa), the Sierra de Perijá (Zulia), and the 

Cordillera de la Costa (Miranda; Map 9). In the park, found 

near La Divisoria de la Concepción, Paramito, and La Aguadita; 

(1,550–)1,700–1,950 m.

This is the smallest Anthurium species found in the park 

apart from A. scandens, which has a very different and distinctive climbing habit.

Anthurium ginesii Croat, in Croat and Lambert, Aroideana 9: 

30, fig. 53. 1986; Morillo and Briceño, in Morillo et al., 

Bot. Ecol. Monocot. Páramos Venez. 1: 69–71, fig. 1. 2011.

FIGURE 23G

Terrestrial or epiphytic herbs; stems to 30 cm long. Leaves simple; cataphylls persisting almost intact or disintegrating into strips 

(not mesh); petioles 25–79 cm long, terete, somewhat flattened 

apically; leaf blades obpyriform, slightly to strongly constricted 

between basal lobes and apex, (14–)20–44 × (6–)10–35 cm, bases 

cordate, margins notably concave, apices gradually to abruptly 

acuminate, basal sinus parabolic to broadly parabolic, 5–15 cm 

deep, lobes divergent, never overlapping, collective vein 2–6 mm 

from margin, subcoriaceous. Inflorescences erect. Peduncles 25–56 

cm long, shorter than leaves. Spathe ovate-lanceolate, (3.5–)8–16 × 

(1.5–)4–5 cm, erect, ± cucullate, green with maroon stripes within, 

uniformly green without. Spadix stipitate, (3.5–)8–16 cm long, ± 

equal to spathe, attenuate-terete, brownish-green to green.

Restricted to South America (Colombia and Venezuela). 

Venezuelan records are from the Andes (Lara, Mérida, Portuguesa, Táchira, and Trujillo) and the Sierra de Perijá (Zulia; Map 

10). In the park, widely distributed in the understory of cloud 

forest; 1,700–2,400 m.

Although Anthurium ginesii and A. nubicola usually can be 

distinguished by their leaf morphology (see key), the two species 

have been confused. This confusion appears to derive from the protologue of the former name, which includes two different elements. 

The type (Croat 54927) and one of the photographs (Croat and 

Lambert, 1986: fig. 53) represent A. ginesii, a species with slightly 

to strongly constricted leaf blades with parabolic to broadly parabolic sinuses. However, at least one of the paratypes (Stergios 

6564), incidentally collected in Guaramacal, and the other photographs (Croat and Lambert, 1986: figs. 51, 52, 56) cited in the protologue clearly represent A. nubicola, which has triangular-ovate 

leaf blades with hippocrepiform to spathulate sinuses.

♦ Anthurium julianii G. S. Bunting, Acta Bot. Venez. 10: 272. 

1975; Morillo and Briceño, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 1: 71–72, fotos 2, 3. 2011.

Epiphytic or hemiepiphytic herbs; stems to 3 m long. Leaves 

simple; cataphylls persisting as a network of fibers; petioles 40–88 

cm long, subterete or narrowly sulcate, equal to or longer than leaf 

blades; leaf blades ovate, 43–71(–75) × 24–46 cm, bases cordate, 

margins slightly undulate, apices acuminate to cuspidate, basal 

lobes rounded, sinus hippocrepiform to spathulate, 12–20 cm deep, 

primary lateral veins 8–12 per side, basal veins 4 or 5 per side, first 

and often second basal veins free, others coalesced 1–3 cm along 

sinus, primary and secondary veins slightly raised above, primary 

and secondary veins prominent below, surface ± rugose, collective 

vein in the apical part of the blade 2–3 mm distant from leaf margin, subcoriaceous. Inflorescences erect or ascending. Peduncles 

14–17.5(–36) cm long. Spathe broadly lanceolate, 10–15(–30) × 

2–3.5(–7) cm, bases clasping, apices caudate, bright or brick-red 

(less commonly reddish-pink) within and without, recurved, persistent. Spadix stipitate, stipe 0.5–2.5 cm, slightly tapered apically, 

10–19(–32) × 1.2–1.8 cm, expanding in fruit, green (at anthesis).

Endemic to the Andes of Venezuela (Lara, Mérida, Táchira, 

and Trujillo; Map 11). In the park, found in montane forest near 

MAP 10. Anthurium ginesii occurrence in Venezuela. MAP 11. Anthurium julianii occurrence in Venezuela.
NUMBER 100 • 1 5

the Laguna de Aguas Negras, in cloud forest on the Fila de Agua 

Fría, and elsewhere on the north slope of Guaramacal; 1,800–

2,900 m.

This species is similar to Anthurium smaragdinum G. S. 

Bunting; characters to distinguish the two are given in the key 

to the genus and following the description of the latter species.

A report by Croat and Carlsen in Hokche et al. (2008) of 

Anthurium julianii occurring in the Cordillera de la Costa (Aragua) needs to be verified.

Anthurium longegeniculatum Engl., Bot. Jahrb. Syst. 25: 379. 

1898; Morillo and Briceño, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 1: 72, fig. 2. 2011.

Terrestrial or epiphytic herbs; stems scandent, to ~1 m 

long. Leaves simple; cataphylls persisting as a braided network 

of fibers covering basal nodes; petioles (16–)30–39 cm long, 

much longer than leaf blades; leaf blades ovate to oblongovate, (9–)13–19.5 × (3–)8–8.5 cm, bases slightly cordate, apices long-acuminate, primary lateral veins ascending, collective 

vein sinuate, loop connected at primary lateral veins, 2–10 mm 

from margin, noticeably closer at apex than base, subcoriaceous, 

brownish glandular-punctate above and below (punctations 

often difficult to see). Inflorescences erect, shorter than petioles. 

Peduncles 15–19.5 cm long. Spathe linear-lanceolate, ~2–2.5 × 

0.8 cm, reflexed, green to greenish-white. Spadix distinctly stipitate, cylindrical, 2–3.5(–7) cm long, green.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, found only in the Cordillera de Mérida 

(Trujillo; Map 12). In the park, found in montane forest on 

the north slope including Qda. Segovia and below Páramo del 

Pumar; 1,800–2,800(–3,000) m.

Anthurium nubicola G. S. Bunting, Acta Bot. Venez. 10: 274. 

1975; Morillo and Briceño, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 1: 73, foto 4. 2011.

Anthurium sp.; Ortega et al., BioLlania 5: 33. 1987 (as to Stergios 2561).

Epiphytic or climbing herbs; stems 0.8–3 m long. Leaves 

simple; cataphylls persisting, degenerating at the base into a fibrous mass; petioles to 60 cm long; leaf blades triangular-ovate, 

not constricted between basal lobes and apex, 28–37(–52) × 11–

15(–28) cm, bases strongly cordate, margins undulate, slightly 

revolute, apices acute to acuminate, basal sinus hippocrepiform 

to spathulate, 8–14 cm deep, lobes not divergent, sometimes 

overlapping, collective nerve present in the upper part of the leaf 

blade, 5–7 mm from margin, coriaceous. Inflorescences erect or 

ascending. Peduncles 25–30 cm long. Spathe 6.5–8(–13) × 1.5–

4(–6.5) cm, green with maroon nerves inside, green outside, ± 

erect. Spadix shortly stipitate, 5–11(–20) cm long, purple.

Endemic to the Cordillera de Mérida of Venezuela (Mérida 

and Trujillo; Map 13). In the park, found in cloud forest on both 

slopes of Guaramacal, including the Fila de Agua Fría; 2,000–

2,900 m.

It appears that Anthurium nubicola is misplaced in A. sect. 

Calomystrium Schott. Although the cataphylls were described in 

the protologue as “persistent, not soon deteriorating,” material 

from Guaramacal that matches the type in leaf morphology has 

cataphylls that disintegrate, at least below, into a mass of fibers.

Anthurium nymphaeifolium K. Koch & C. D. Bouché, Index 

Seminum [Berlin] 1853: 16. 1853 (“nymphaefolium”); Morillo and Briceño, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 1: 73–74, fotos 5, 10. 2011.

FIGURE 23C, D

Anthurium sp.; Ortega et al., BioLlania 5: 33. 1987 (as to Stergios 2547).

MAP 12. Anthurium longegeniculatum occurrence in Venezuela.

MAP 13. Anthurium nubicola occurrence in Venezuela.
1 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Terrestrial, epiphytic or climbing herbs; stems elongate, to 

2 m long. Leaves simple; cataphylls reddish-brown, entire, and 

persistent; petioles 50–83 cm long, erect, rigid, ± terete; blades 

ovate to broadly ovate, 42–51 × 24–28 cm, lateral veins prominent, joined near the apex of the blade to form a collective vein, 

basal sinus hippocrepiform to spathulate, 9–13 cm deep, basal 

lobes ± as long as wide, sometimes almost touching, coriaceous, 

nitid. Inflorescences erect. Peduncles 35–76 cm long, equal to 

or longer than petioles. Spathe ovate to broadly ovate, 10–17 

× 4–6.5 cm, erect, apices cuspidate or long-acuminate, white, 

pinkish-white or greenish-white within, green without. Spadix 

stipitate, cylindrical, 5.5–12 × ~1.5 cm, shorter than spathe.

Found in South America (Colombia and Venezuela). Widespread in Venezuela (Amazonas, Anzoátegui, Aragua, Barinas, 

Bolívar, Carabobo, Cojedes, Distrito Federal, Falcón, Lara, 

Mérida, Miranda, Monagas, Portuguesa, Táchira, Trujillo, Yaracuy, and Zulia; Map 14). Found throughout the park in montane forest and cloud forest; (1,500–)1,800–2,900 m.

This is a variable species for which several varieties have 

been described, although none are recognized here. Bunting 

(1979) attempted to correlate some of the morphological variation (spathe color, leaf shape, etc.) that he observed with geography and elevation but stopped well short of proposing a formal 

subspecific taxonomy. The limits of Anthurium nymphaeifolium

and related species assigned to A. sect. Calomystrium, including 

A. ramoncaracasii, deserve further study.

♦ Anthurium ramoncaracasii Stergios & Dorr, Acta Bot. Venez. 

27: 96, fig. 1. 2004 [2005].

FIGURES 2, 23E

Terrestrial herbs; stems scandent, to 2 m long. Leaves simple; cataphylls persistent, entire; petioles (50–)73–80 cm long, 

subterete, sulcate near base; leaf blades ovate to broadly ovate, 

(20–)50–65 cm × 25–35 cm wide, bases cordate, margins slightly 

sinuous, apices acute to acuminate, basal sinus parabolic to spathulate, 8–13 cm deep, basal lobes rounded, primary lateral veins 

6–12 per side, basal veins 4–5 per side, all ± free, midvein and 

basal nerves prominent below, collective vein originating from uppermost basal vein, extending length of apical lobe, 5–12 mm from 

margin, subcoriaceous. Inflorescences ± pendent. Peduncles 45–68 

cm long. Spathe broadly ovate, cymbiform, 10–18 × 5–8 cm, 

abruptly acuminate, slightly cucullate, light greenish-cream to 

greenish-yellow, persistent. Spadix stipitate, cylindrical, slightly 

tapering apically, 14–21 × 1.5–1.8 cm, arcuate, declined 45°–90°, 

bright wine-red in color, longer than the spathe (especially in fruit).

Endemic to the Cordillera de Mérida in Venezuela (Portuguesa and Trujillo; Map 15). In the park, found on the north 

slope of Guaramacal; 1,750–2,600 m.

This species is very similar to Anthurium nymphaeifolium,

from which it can be distinguished by its larger leaf blades, 

greenish-cream to greenish-yellow spathe, and spadix that exceeds the spathe in length.

Anthurium scandens (Aubl.) Engl., in Martius, Fl. Bras. 3(2): 78. 

1878; Bunting, in Berry et al., Fl. Venez. Guayana 2: 609, 

fig. 545. 1995; Morillo and Briceño, in Morillo et al., Bot. 

Ecol. Monocot. Páramos Venez. 1: 76–77, foto 6. 2011. 

Dracontium scandens Aubl., Hist. Pl. Guiane 2: 836. 1775.

Only the nominate subspecies is found in Guaramacal.

Anthurium scandens subsp. scandens

Epiphytic or climbing herbs; stems <1 m long, scandent or 

firmly attached to substrate. Leaves simple; cataphylls persisting 

as a thin lattice or network of fibers around the stems; petioles 

MAP 14. Anthurium nymphaeifolium occurrence in Venezuela. MAP 15. Anthurium ramoncaracasii occurrence in Venezuela.
NUMBER 100 • 1 7

FIGURE 2. Araceae, Anthurium ramoncaracasii Stergios & Dorr. A. Leaf blade. B. Section of stem showing one node and cataphylls. C. Spadix and spathe at anthesis. D. Detail of spadix showing mature flowers. E. Flower (longitudinal view). F. Flower (apical view). G. Tepal with 

reflexed tip. H. Flower (longitudinal view), tepals removed to show stamens. I. Flower (longitudinal section). J. Immature seed. (A–J, Stergios 

et al. 20101.)
1 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

1.2–3(–5.5) cm long, shorter than leaf blades; leaf blades ovate 

to ovate-lanceolate, 4.5–10 × 1.2–3(–5.5) cm, bases cuneate, apices acute to acuminate, primary lateral veins pinnate-parallel, 

ascending, collective vein arising from the base and extending 

to the apex, 2–3 mm from the margin, subcoriaceous, blackish 

glandular punctations above and below. Inflorescences often exceeding petioles in length. Peduncles 1.4–3(–4.5) cm long. Spathe 

lanceolate to oblong-lanceolate, 0.7–1 × 0.2 cm, recurved, light 

green. Spadix sessile or short-stipitate, cylindrical to slightly 

attenuate, 0.7–1.5 cm long (expanding in fruit), whitish- or 

grayish-green. Berries white or pale lavender.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Trinidad and Tobago, 

the Guianas, Ecuador, Peru, Bolivia, and Brazil). Widespread 

in Venezuela (Amazonas, Anzoátegui, Apure, Aragua, Barinas, 

Bolívar, Carabobo, Delta Amacuro, Distrito Federal, Falcón, 

Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, 

Sucre, Táchira, Trujillo, Yaracuy, and Zulia; Map 16). Found on 

both slopes of Guaramacal; 1,650–2,300(–3,000) m.

The stems and leaves of Anthurium scandens subsp. scandens attach themselves tightly to tree trunks, and this alone distinguishes this taxon from all other taxa of Anthurium found in the 

park. In addition, A. scandens subsp. scandens has the smallest 

leaves and inflorescences of all of the species of Anthurium known 

to occur in Guaramacal. The white or pale lavender berries are 

often the most conspicuous part of the very small inflorescences.

Anthurium scandens evidently is a polyploid complex (Sheffer et al., 1980), but generally, correlations between ploidy level 

and morphology have not been established. A diploid found in 

Costa Rica was segregated as A. scanden subsp. pusillum R. 

Sheffer. Morphologically, it can be distinguished from the nominate subspecies by its smaller (0.2–1 vs. 0.7–1.5 cm long) spadix 

with fewer (5–12 vs. 16–60) flowers.

Anthurium smaragdinum G. S. Bunting, Acta Bot. Venez. 10: 

278. 1975.

FIGURE 23F

Anthurium humboldtianum subsp. viridispadix Croat, in Croat and Lambert, Aroideana 9: 36, figs. 64–66. 1986.

Anthurium subsagittatum auct., non (Kunth) Kunth; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 38. 2000 [2001].

Epiphytic or climbing herbs; stems to 8 m long. Leaves simple; cataphylls persisting as a network of fibers; petioles 50–61 

cm long, terete to broadly sulcate, ± equal to blade in length; 

leaf blades ovate to ovate-triangular, 48–61 × 26–38 cm, bases 

cordate, margins undulate, apices acute to acuminate, basal sinus 

hippocrepiform, 9–12 cm deep, basal lobes rounded, primary 

veins scarcely raised above, primary and secondary veins prominent below, surface ± rugose, collective vein in the apical part of 

the blade 5–10 mm from the blade margin, subcoriaceous. Inflorescences erect. Peduncles 32–53 cm long. Spathe lanceolate to 

narrowly lanceolate, 12–21 × 1.5–3 cm, base clasping, apex longacuminate, green, spreading, recurved at anthesis. Spadix stipitate, inserted obliquely, narrow, slightly tapered apically, 12–21 × 

0.6–0.8 cm (in flower), 21–28 × 1–1.2 cm (in fruit), green.

Restricted to South America (Colombia and Venezuela). 

In Venezuela, found in the Andes (Lara, Mérida, Portuguesa, 

Táchira, and Trujillo) and possibly the Sierra de Perijá (Zulia; 

Map 17). In the park, found in montane forest and cloud forest 

on both slopes of Guaramacal; 1,600–2,600 m.

MAP 16. Anthurium scandens subsp. scandens occurrence in 

Venezuela. MAP 17. Anthurium smaragdinum occurrence in Venezuela.
NUMBER 100 • 1 9

This species is similar to Anthurium julianii and shares the 

same habitat, but it can be distinguished by spathe color (completely green vs. red) and the distance from which the collective 

vein is found from the leaf margin (5–10 vs. 2–3 mm).

Monstera Adans.

Monstera Adans., Fam. Pl. 2: 470. 1763, nom. cons.

Epiphytic, hemiepiphytic or scandent herbs; laticifers absent; trichosclereids abundant. Leaves distichous; sheaths long, 

persistent or rotting and forming a fibrous mass or entirely deciduous; petioles geniculate apically, pulvinus present at base 

and apex; leaf blades simple, oblong to ovate-elliptic, often 

oblique, margins entire, blade perforate or not, when perforate 

perforations sometimes extending to margins of blade, apices 

various, primary lateral nerves pinnate, running into marginal 

nerves, secondary lateral nerves parallel to the primary ones 

or reticulate, membranous to subcoriaceous. Inflorescences 1 

to several. Peduncles terete or flattened, shorter than petioles. 

Spathe ovate or oblong-ovate, usually erect, cuspidate, hemispherical to naviculiform, base ± convolute, white to lilac-pink 

within, often yellow when mature, deciduous. Spadix sessile, 

subcylindrical, shorter than spathe, sterile near the base. Fertile flowers: bisexual, perigone absent; stamens 4, free; filaments flattened; gynoecium obovoid to ellipsoid, prismatic; 

ovary 2-locular; ovules 2 per locule, anatropous; stylar region 

broader than ovary, apex truncate to shortly attenuate; stigma 

oblong-elliptic to linear and longitudinal or round. Sterile flowers: staminodia 4, minute; pistillode 2-locular, prismatic; ovules 

0. Berry 1–3-seeded, pulp gray or orange. Seeds globose; testa 

soft, smooth.

A neotropical genus of 20–40 species found in Central 

America, the West Indies, and South America (Colombia, Venezuela, Curaçao, Trinidad and Tobago, the Guianas, Ecuador, 

Peru, Bolivia, Brazil, Paraguay, and Argentina). Seven species 

occur in Venezuela.

References. Engler and Krause (1908); Madison 

(1977); Tam et al. (2004).

The leaves of Monstera are heteroblastic; juvenile and adult 

leaves can be strikingly different. Adult leaves are as described 

above, whereas juvenile leaves have very short petioles and their 

blades are held fast against their substrate in an overlapping, 

shingle-like pattern.

Monstera has not been sampled extensively for molecular 

analyses, but the scant data that exist suggest that the genus is 

monophyletic and part of a clade (Rhaphidophora clade) consisting of six paleotropical genera within which Monstera forms 

a subclade with the southeast Asian genera Amydrium Schott 

and Epipremnum Schott (Tam et al., 2004; Cusimano et al., 

2011). Rhaphidophora Hassk. has a more extensive distribution 

than Monstera and is found not only in southeast Asia but also 

in Australia and West Africa. Current molecular data generally 

support the evolutionary relationships of Monstera that Madison (1977) inferred on the basis of morphology.

Monstera adansonii Schott, Wiener Z. Kunst 1830: 1028. 1830. 

Dracontium pertusum L., Sp. Pl. 968. 1753, non Monstera 

pertusa (Roxb.) Schott, Wiener Z. Kunst 1830: 1028. 1830.

Monstera adansonii comprises three varieties found in Central America, the West Indies, and South America (Colombia, 

Venezuela, Curaçao, Trinidad and Tobago, the Guianas, Ecuador, 

Peru, Bolivia, and Brazil). The nominate variety is restricted to the 

West Indies, whereas the other two occur in Venezuela. The following variety is the only one known from the Venezuelan Andes.

Monstera adansonii var. laniata (Schott) Madison, Contr. Gray 

Herb. 207: 38. 1977; Bunting, in Berry et al., Fl. Venez. 

Guayana 2: 628. 1995. Tornelia laniata Schott, Oesterr. Bot. 

Z. 8: 179. 1858.

Adult stems elliptic in cross section. Leaves: petioles (20–)

40–45 cm long; sheath wings neatly deciduous; leaf blades 

ovate to broadly ovate, 15–40 × 20–55 cm (length less than 

2× width), bases unequal, one side truncate to subcordate, the 

other side cuneate to acute, margins entire, perforations absent 

to many, apices acute to shortly mucronate, central nerve and 

lateral principal nerves very light in color with respect to the 

rest of the blade, pale color diminishing toward the apex, subcoriaceous. Peduncles 15–20 cm long, subterete. Mature spathe 

cream-colored to yellowish, 12–28 × 10–25 cm, ~2× longer than 

the spadix, coriaceous. Flowering spadix yellowish, cylindrical, 

10–12 × 2–3 cm; gynoecium prismatic, truncate apically.

Found in Central America, the West Indies (Grenada), and 

South America (Colombia, Venezuela, Curaçao, Trinidad and 

Tobago, the Guianas, Ecuador, Peru, Bolivia, and Brazil). Widespread in Venezuela (Amazonas, Anzoátegui, Apure, Aragua, 

Barinas, Bolívar, Carabobo, Delta Amacuro, Distrito Federal, 

MAP 18. Monstera adansonii var. laniata occurrence in Venezuela.
2 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Falcón, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, 

Táchira, Trujillo, Yaracuy, and Zulia; Map 18). Found in forest 

in the karst area above the Río Amarillo near the lower limits of 

the southern boundary of the park; 1,200–1,400 m.

Our variety can be distinguished from Monstera adansonii

var. klotzschiana (Schott) Madison, the only other variety found 

in Venezuela, by its leaves. Sheath wings are deciduous and 

blades are less than 2× as long as wide in our variety, whereas 

sheath wings are persistent and blades more than 2× as long as 

wide in M. adansonii var. klotzschiana.

Philodendron Schott

Philodendron Schott, Wiener Z. Kunst 1829: 780. 1829, nom. & orth. cons.

Epiphytic, hemiepiphytic, climbing, vine-like or terrestrial 

herbs; stems repent to rhizomatous, sometimes producing flagelliform shoots; laticifers present. Leaves numerous, small to 

very large; prophylls of mature stems caducous, marcescent and 

deciduous, or persistent and rotting to form a fibrous mass; 

petioles sometimes warty or covered with a scaly indumentum, 

sometimes swollen, rarely geniculate apically; sheaths elongate 

and slightly ligulate in monopodial leaves or very short and inconspicuous except when subtending inflorescences; leaf blades 

variable in shape, simple and linear, cordate, sagittate or hastate, or variously dissected, resin canals visible below, primary 

lateral veins pinnate, forming 1 marginal vein, higher-order venation parallel-pinnate. Flowering branches conforming to 1 of 

3 general sympodial patterns: prophyll, many foliage leaves, 1 

or 2(3) inflorescences (subgen. Pteromischum (Schott) Mayo); 

prophyll, subsequent internode suppressed, 1 foliage leaf, 1–11 

inflorescences (subgen. Philodendron); or prophyll, subsequent 

internode developed or not, 1 foliage leaf, 1(–2) inflorescences 

(subgen. Meconostigma (Schott) Engler). Inflorescences 1–11 

per floral sympodium, secreting at anthesis a resinous substance 

from the spathe or spadix but rarely from both. Peduncles usually shorter than petioles. Spathe erect, persistent, deciduous at 

base when fruit matures, thickened, usually constricted between 

blade and tube, tube convolute, cylindrical, often reddish-purple 

within, blade naviculiform, widely ringent at anthesis then closing, usually white within. Spadix sessile to stipitate, pistillate 

zone free, shorter than the staminate zone and separated by a 

sterile zone of staminodial flowers, which is thicker in diameter 

than staminate zone. Flowers unisexual; perigone absent. Staminate flowers: 2–6-merous; stamens free; anthers sessile or almost 

sessile. Sterile staminate flowers: staminodes usually prismatic. 

Pistillate flowers: gynoecium ovoid, subcylindrical or obovoid; 

ovary usually 4–8-locular; ovules 1–50(or more) per locule, 

usually hemiorthotropous; stylar region as broad as ovary or 

broader; stigma sometimes lobed or discoid-hemispheric. Berry 

subcylindrical to obovoid, 1- to many-seeded, white, whitish 

translucent, red or orange-red. Seeds ovoid-oblong to ellipsoid, 

variable in size; testa costate.

A neotropical genus of ~750 species found in Mexico, Central America, the West Indies, and South America (Colombia, 

Venezuela, Trinidad and Tobago, the Guianas, Ecuador, Peru, 

Bolivia, Brazil, Paraguay, and Argentina); introduced in Africa, 

Asia, Australia, and the Pacific Islands. In Venezuela, ~108 species have been recorded.

References. Gauthier et al. (2008); Grayum 

(1996); Krause (1913).

The relationship between Philodendron and Homalomena

Schott, a genus found in tropical America and Asia, is unresolved 

(Cabrera et al., 2008; Gauthier et al., 2008). Nonetheless, these 

two genera and several smaller ones form a strongly supported 

clade (Philodendron clade) that is united synapomorphically by 

a pistillate-sterile-staminate pattern of spadix floral zonation 

(Cusimano et al., 2011).

The three subgenera of Philodendron (see generic description above) originally recognized on morphological criteria alone 

are supported as monophyletic by molecular data (Gauthier et 

al., 2008). However, this classification could change with resolution of the larger problem concerning generic boundaries.

KEY TO THE SPECIES OF PHILODENDRON

1a. One foliage leaf per sympodial branch (subgen. Philodendron); leaf blades cordate-oblong or cordate-ovate, 25–45 cm 

long; petioles sheathed at base only . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. fraternum

1b. Many foliage leaves per sympodial branch (subgen. Pteromischum); leaf blades broadly lanceolate or narrowly ovate to 

ovate, (9.5–)12–14 cm long; petioles completely surrounded by a winged sheath . . . . . . . . . . . . . . . . . . P. tuerckheimii

Philodendron fraternum Schott, Bonplandia (Hannover) 7: 29. 

1859.

FIGURE 23H

Hemiepiphytic or scandent liana-like herbs. Leaves simple; 

petioles 30–35 cm long, sheaths at base only, to 7 cm long; leaf 

blades cordate-oblong or cordate-ovate, 25–45 × 10–15(–25) 

cm, basal lobes 3–12 × 5–7 cm, sinus to 12 cm deep, parabolic 

to subhippocrepiform, margins entire, apices acuminate to longcuspidate. Inflorescences 2 or 3 per axil. Peduncles 4–7 cm long. 

Spathe 6–7 cm long, cream to greenish-white within, maroonpurple without (in bud), becoming green to creamy yellow (at 

anthesis). Spadix ~7 cm long, white.

Found in South America (Colombia and Venezuela). 

Widespread in Venezuela (Apure, Aragua, Distrito Federal, 

Falcón, Lara, Mérida, Miranda, Portuguesa, Táchira, Trujillo, 

Yaracuy, and Zulia; Map 19). In the park, found in montane 

and cloud forest on both slopes of Guaramacal; 1,800–2,350(–

2,750) m.
NUMBER 100 • 2 1

Philodendron tuerckheimii Grayum, Syst. Bot. Monogr. 47: 

174, figs. 105–108. 1996.

Philodendron rudgeanum auct., non Schott; Ortega et al., BioLlania 5: 33. 

1987.

Greatly branched, scandent hemiepiphytic or vine-like herbs. 

Leaves simple; petioles (3.5–)5–6 cm long, completely surrounded 

by a winged sheath; leaf blades broadly lanceolate or narrowly 

ovate to ovate, (9.5–)12–14 × 4–5 cm, bases cuneate to rounded 

or truncate, margins entire, apices subfalcate to acuminate. Inflorescences solitary. Peduncles 1–3 cm long. Spathe 7–14 × 1–3 cm, 

pale green to whitish within, cream to greenish-white or yellowgreen without. Spadix 7–12 cm long, white or creamy white.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, and Ecuador). In Venezuela, found in the 

Andes (Lara, Mérida, Portuguesa, Táchira, and Trujillo) and the 

Cordillera de la Costa (Carabobo and Yaracuy; Map 20). In the 

park, found in montane forest on the south slope of Guaramacal; 

(1,600–)1,800–1,900 m.

Grayum (1996) acknowledges that this wide-ranging species is not well defined and might eventually prove divisible into 

two or more species. He also notes that South American material 

differs from Mexican and Central American material in having 

more narrowly winged petiole sheaths, longer and proportionally narrower leaf blades, and longer spadices.

Rhodospatha Poepp.

Rhodospatha Poepp., in Poeppig and Endlicher, Nov. Gen. Sp. Pl. 3: 91. 

1845.

Scandent or suberect herbs, often with flagelliform shoots; laticifers absent; trichosclereids abundant. Leaves simple, distichous; 

petioles elongate, bases encircling stem, apices geniculate; sheaths 

persistent to marcescent; leaf blades oblong or elliptic, ± oblique, 

margins entire, primary lateral nerves pinnate, running into a ± distinct marginal nerve, secondary and tertiary lateral nerves parallelpinnate. Inflorescences usually solitary. Peduncles shorter to longer 

than petioles. Spathe large, broadly ovate or oblong-ovate, cuspidate, yellowish-white, cream or purplish within, scarcely opening 

at anthesis, soon decaying or caducous. Spadix erect, long-stipitate 

to sessile, cylindrical-conical. Flowers bisexual or basalmost ones 

sterile or pistillate and scattered, perigone absent. Stamens 4, free, 

filaments flattened. Gynoecium compressed, obconical to cylindrical; ovary 2-locular; ovules numerous per locule, anatropous to 

hemianatropous; stylar region well developed, broader than ovary, 

prismatic, truncate to convex apically; stigma elliptic to linear. 

Berry cylindrical-prismatic, truncate, few- to many-seeded. Seeds 

lenticular; testa smooth or verrucose.

A neotropical genus of ~75 species (many undescribed) 

found in Mexico, Central America, and South America (Colombia, Venezuela, Trinidad and Tobago, the Guianas, Ecuador, 

Peru, Bolivia, and Brazil). A dozen species occur in Venezuela.

References. Engler and Krause (1908); Tam et al. 

(2004).

Molecular data (Cabrera et al., 2008; Cusimano et al., 2011) 

place Rhodospatha in an exclusively neotropical clade (Heteropsis

clade), which also includes Alloschemone Schott, Heteropsis Kunth, 

and Stenospermation Schott. This clade, the Rhaphidophora clade 

(see Monstera above), and the Spathiphylleae (Spathiphyllum Schott 

and Holochlamys Engl.) constitute the Monsteroideae, which is sister to the Pothoideae. Cusimano et al. (2011) call the clade that 

includes these two subfamilies the bisexual climbers clade.

MAP 19. Philodendron fraternum occurrence in Venezuela.

MAP 20. Philodendron tuerckheimii occurrence in Venezuela.
2 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Rhodospatha badilloi G. S. Bunting, Acta Bot. Venez. 10: 318. 

1975.

Rhodospatha latifolia auct., non Poepp.; Dorr et al., Contr. U.S. Natl. Herb. 

40: 38. 2000 [2001].

Scandent herbs; aerial roots arising from basal nodes. 

Leaves: cataphylls linear-lanceolate, 3–4 cm long; petioles 

shorter than or equal to the leaf blade, petiolar wings wavy basally but straight and attenuate approaching the geniculum; leaf 

blades ovate-elliptic, 37–39 × 16–17 cm, entire, cuspidate, primary and secondary nerves impressed above, noticeably raised 

below where the principal secondary nerves are uniformly parallel on both sides of the midnerve, membranous to semicoriaceous. Spathe and spathe bracts deciduous (not seen). Spadix 

20–21(–30) × 2–3 cm; gynoecium asymmetrical.

Endemic to Venezuela, where it is known from the Andes 

(Lara, Táchira, and Trujillo) and the Cordillera de la Costa (Aragua; Map 21). In the park, found in forest near La Punta of El 

Santuario; 1,900 m.

Xanthosoma Schott

Xanthosoma Schott, in Schott and Endlicher, Melet. Bot. 19. 1832.

Small to gigantic, sometimes arborescent, terrestrial herbs; 

rhizomatous or with corms; stems erect or acaulescent; latex 

milky or watery. Leaves glabrous, rarely pubescent, sometimes 

aggregated at the apex of aerial rhizomes (our species); blades 

entire (our species) or divided into 3 to many segments, cordate, 

sagittate or hastate; basal ribs well developed, often naked proximally; primary lateral nerves pinnate and forming a submarginal 

collective nerve; petioles sheathed below. Inflorescences 1 to 

several, always appearing with leaves. Peduncles usually shorter 

than petioles. Spathe strongly constricted, tube ovoid to ellipsoid, green, red or purplish, persistent, expanded blade naviculiform to oblong-lanceolate, ivory or pink to purplish, marcescent 

after anthesis and then deciduous. Spadix shorter than spathe; 

pistillate zone short, separated from staminate zone by sterile 

staminate flowers; staminate zone longer than pistillate zone 

and fertile to apex. Flowers unisexual, perigone absent. Staminate flowers: 4–6-merous, stamens united in a truncate synandrium. Pistillate flowers: ovary ovoid, (1–)2–4-locular; ovules 

(12–)20 or more, anatropous or hemianatropous; stylar region 

broader than ovary; stigma discoid or 2–4-lobed. Berry cylindric, ± furrowed apically, whitish to orange, many-seeded. Seeds 

ovoid, small; testa costate.

A neotropical genus of ~60 species found in Mexico, Central America, the West Indies, and South America (Colombia, 

Venezuela, Trinidad and Tobago, the Guianas, Ecuador, Peru, 

Brazil, Paraguay, and Argentina). Approximately 22 species are 

found in Venezuela.

References. Engler and Krause (1920); Gonçalves 

(2011).

Cusimano et al. (2011) placed Xanthosoma in a clade (tribe 

Caladieae) that they argue is also supported by several anatomical and morphological synapomorphies, including the presence 

of anastomosing laticifers and a persistent spathe tube with a 

deciduous or marcescent spathe blade.

A number of Xanthosoma species are economically important because their corms are used as food. Nonetheless, no 

comprehensive revision of the genus exists. Gonçalves (2011) 

enumerated some of the reasons why the taxonomy of Xanthosoma is chaotic and unsatisfactory: (1) Relatively few herbarium 

specimens are available for study, and those that exist are often 

poorly prepared. (2) Many species names were based on incomplete material, and the application of these names is a matter of 

debate. (3) Most species are phenotypically plastic. (4) There 

has been a great amount of human-mediated selection influencing the morphology and distribution of species that are used 

as food.

Xanthosoma sagittifolium (L.) Schott, in Schott and Endlicher, 

Melet. Bot. 19. 1832 (“sagittaefolium”); Bunting, in Berry 

et al., Fl. Venez. Guayana 2: 677. 1995. Arum sagittifolium

L., Sp. Pl. 966. 1753 (“sagittaefolium”).

FIGURE 23I, J

Very large, erect herbs, (0.5–)1–3.5 m tall; rhizomes 0.5–

2.5 m tall; latex watery, light yellow or whitish (very irritating 

to the skin). Leaves 4–7 arising from the apex of the rhizome; 

petioles 1–1.6(–2.5) m long, green, mottled or not mottled; leaf 

blades broadly ovate, (0.6–)1.2–1.5 × 0.9–1.1 m, reflexed, basal 

lobes mostly overlapping, with a closed or narrowly obovate 

sinus, margins slightly undulate, apices obtuse, membranous. 

Inflorescences 1–3 per axil. Spathe erect, 20–30(–40) cm long, 

green; tube 13 × 4.7 cm (at base) or 2 cm (at constriction), green 

MAP 21. Rhodospatha badilloi occurrence in Venezuela.
NUMBER 100 • 2 3

outside and inside or green outside and weakly to heavily tinged 

purple inside or dark purple outside and inside; blade erect, 

ovate-elliptic, somewhat cucullate, ivory. Spadix sessile, 15–25

(–30) cm long, white, creamy yellow below.

Found in Central America, the West Indies, and South 

America (Colombia, Venezuela, Trinidad and Tobago, Suriname, 

French Guiana, Ecuador, Peru, Bolivia, and Brazil); introduced 

and/or cultivated in North America (USA), Africa, and Asia. 

Widespread in Venezuela (Amazonas, Barinas, Delta Amacuro, 

Mérida, Miranda, Nueva Esparta, Portuguesa, Sucre, Táchira, 

Trujillo, and Yaracuy; Map 22). In the park, found in forest understory on the south slope near Qda. Honda and La Punta and 

along the Boconó–Guaramacal road; 1,800–1,900(–2,700) m.

Common Names. Guaje, guaje bravo, and guaje 

simangue.

The application of the name Xanthosoma sagittifolium is 

problematic. We employ the name here in the sense that it generally has been used in Venezuela (see, e.g., Bunting, 1979; Bunting 

in Berry et al., 1995). Gonçalves (2011), however, argues that the 

name often is misapplied, and he restricts the native range of X. 

sagittifolium to the West Indies (the type locality) and argues that 

the species is rarely cultivated elsewhere. This may be correct, 

but our material still keys to X. sagittifolium in his synopsis of 

the commonly cultivated species.

Guaramacal material shows remarkable variation in spathe 

tube color and in the presence or absence of maculations on 

petioles and peduncles (none of these differences are seen in herbarium material). According to Croat (in litt.), spathe tube color 

generally is a reliable character in Xanthosoma, but in our material it ranges from green inside and outside to weakly to heavily 

tinged purple inside. The maculations on petioles and peduncles 

also are inconstant in the Guaramacal populations and, as far as 

we can tell, are not correlated with any other character. These 

maculations have not been previously noted in descriptions of X. 

sagittifolium. Irrespective of these color differences, plants found 

in Guaramacal are relatively uniform in terms of habit and leaf 

blade shape and size.

Xanthosoma sagittifolium is not currently used as food in 

Guaramacal (R. Caracas, pers. comm.), but that does not exclude the possibility that it was introduced and used as food in 

the past.

Xanthosoma pubescens Poepp. also is found in Portuguesa 

near the park but at much lower elevations (900–1,200 m). It 

can be distinguished from X. sagittifolium by its pilose foliage 

and much smaller size.

Arecaceae

L. J. DORR AND S. MIGUEL NIÑO

Small, medium-sized or large hermaphroditic, polygamous, monoecious or dioecious plants; stems (trunks) woody, 

solitary or clustered, erect, sometimes acaulescent or climbing 

or subterranean; unbranched (our taxa) or rarely branched; 

stilt roots sometimes present; unarmed or spiny. Leaves spirally arranged, rarely distichous or polystichous; leaf sheaths 

tubular, closed or open (due to rupturing); petioles present; leaf 

blades palmate, costapalmate (i.e., with a petiole that extends 

into the leaf blade), pinnate, bipinnate or entire, plicate in bud, 

splitting along the adaxial folds (induplicate) or abaxial folds 

(reduplicate) or not splitting. Inflorescences axillary, solitary 

or multiple, spicate or 1–6-branched, interfoliar or infrafoliar; prophyll 2-keeled; peduncular bract(s) present, rarely absent. Flowers usually 3-merous; bisexual or unisexual, sessile 

or stalked. Sepals (2)3(rarely more), free or connate. Petals 

(2)3(rarely more), free or united. Stamens (3)6(or many). Staminodes present in pistillate flowers. Gynoecium apocarpous or 

syncarpous, sometimes pseudomonomerous, 3(–10)-locular or 

frequently with only 1 fertile locule; style distinct or not; stigmas erect or recurved. Fruits 1(2–10)-seeded, fleshy or dry and 

fibrous, globose or ellipsoid, often hard. Seeds 1(–2 or more); 

endosperm homogeneous or ruminate; embryo apical, lateral 

or basal.

A mostly pantropical family of ~190 genera and ~2,500 species with taxa also found in subtropical and temperate regions of 

North America (USA), South America, Europe, Africa, Asia, and 

Australia. Thirty genera and ~100 species are known to occur in 

Venezuela.

References. Baker and Couvreur (2013); Baker et 

al. (2009, 2011); Braun and Delascio (1987); Dransfield and 

Uhl (1998); Dransfield et al. (2005, 2008); Henderson et al. 

(1995); Stauffer (1999); Tomlinson et al. (2011); Wessels Boer 

(1971).

MAP 22. Xanthosoma sagittifolium occurrence in Venezuela.
2 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

KEY TO THE GENERA OF ARECACEAE

1a. Stems and/or leaves with spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aiphanes

1b. Stems and leaves unarmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Leaf margins dentate; ultimate leaflets (pinnae) flabellate (i.e., fan-shaped), widely toothed . . . . . . . . . . . Wettinia

2b. Leaf margins entire; ultimate leaflets (pinnae) bifid or simple, never flabellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Arborescent plants, to 15 m tall; stems (trunks) 10–20 cm in diameter; ultimate leaflets (pinnae) simple; sheaths 

reddish to purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prestoea

 3b. Shrubby or arborescent plants, 1–8 m tall, stems 0.5–8 cm in diameter; ultimate leaflets (pinnae) bifid; sheaths 

green or dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Dioecious plants; leaflets (pinnae) alternate or almost alternate, sigmoidal; leaf sheaths closed or only open 

apically, green or yellowish-green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chamaedorea

 4b. Monoecious plants; leaflets (pinnae) opposite or subopposite, deltoid; leaf sheaths open, light or dark brown 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geonoma

Aiphanes Willd.

Aiphanes Willd., Samml. Deutsch. Abh. Königl. Akad. Wiss. Berlin 1803: 

250. 1806.

Monoecious, acaulescent or caulescent plants; stems (trunks) 

solitary or clustered, branched or not; spiny. Leaves spirally arranged or distichous; leaf sheaths open, densely spiny; leaf blades 

pinnate, rarely entire, spiny; leaflets (pinnae) with serrate apices 

or irregularly divided. Inflorescences generally 1(2)-branched, 

rarely spicate, interfoliar, erect. Flowers unisexual, 3-merous. 

Staminate flowers: sepals 3, free or shortly connate; petals 3, 

free or shortly connate; stamens 6, in 2 series; filaments basally 

connate. Pistillate flowers: usually larger than staminate flowers; sepals 3, free, cartilaginous; petals 3, connate 1⁄2 their length; 

staminodes 6, fused; gynoecium 3-locular, 3-ovulate; stigmas 

3. Fruits 1-seeded, globose or ellipsoid, red, white, orange or 

purple. Seeds globose; endosperm homogeneous; embryo lateral. 

[Arecoideae: Cocoseae: Bactridinae.]

A neotropical genus of ~25 species found in Central America (Panama), the West Indies, and South America (Colombia, 

Venezuela, Trinidad and Tobago, Ecuador, Peru, Bolivia, and 

Brazil). Two species occur in Venezuela, one widespread and one 

localized in the Andes.

References. Bernal and Borchsenius (2010); Borchsenius and Bernal (1996).

Aiphanes is easily recognized because it is the only genus of 

spiny palm in the Americas with leaflets that have jagged tips.

Aiphanes lindeniana (H. Wendl.) H. Wendl., in Kerchove de 

Denterghem, Palmiers 230. 1878. Martinezia lindeniana H. 

Wendl., Linnaea 28: 349. 1856 [1857].

FIGURE 3

♦ Aiphanes stergiosii M. Niño et al., Sida 21: 1600, fig. 1. 2005.

Aiphanes sp. A; Dorr et al., Contr. U.S. Natl. Herb. 40: 39. 2000 [2001].

Small to medium-sized plants; stems (trunks) solitary (our 

material) or clustered, 4–6 m × 3–4.5 cm, densely spiny, spines 

usually retrorse, 1–9 cm long, brownish-black, clustered in the 

center of internodes. Leaves 4–8(–10) per plant; leaflets (pinnae) 

18–24(–45) per side, inserted in groups of 3–4(–7), lanceolate to 

narrowly cuneate, middle leaflets 10–20(–34) × (1–)5–8.5 cm, 

margins and midvein above and below spiny, apical leaflets almost always divided 2× or more, with an irregular apical margin. 

Inflorescences 1-branched; peduncular bracts 80–90(–140) cm × 

4–5 mm, tomentose with several scattered spines; peduncles 90–

120 cm × 3–5 mm, densely spiny at the base; rachillae densely 

tomentose. Fruits subglobose, 9–15 mm in diameter, red or orange, smooth and glossy.

Found in South America (Colombia and Venezuela). In Venezuela, known at present only from the Andes (Portuguesa; Map 

23). La Divisoria de la Concepción; 1,700–1,900 m.

Common Name. Macanilla.

Our collections of this species are remarkably disjunct from 

the nearest Colombian population, but we suspect this might be 

an artifact that can be attributed to inadequate floristic sampling 

of the east-facing slopes of the Venezuelan Andes.

MAP 23. Aiphanes lindeniana occurrence in Venezuela.
NUMBER 100 • 2 5

FIGURE 3. Arecaceae, Aiphanes lindeniana (H. Wendl.) H. Wendl. A. Leaf apex (most apices are eventually divided as a consequence of environmental factors). B. Medial portion of leaf. C. Leaf base. D. Staminate and pistillate flowers. E. Staminate flowers. F. Fruit. G–I. Anthers. 

J. Pistillode. K, L. Sepals (staminate flower). (Niño & Stergios 1431.)
2 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Chamaedorea Willd.

Chamaedorea Willd., Sp. Pl. 4(2): 638, 800. 1806, nom. cons.

Dioecious, shrubby or sometimes arborescent plants; stems 

(trunks) solitary or clustered, procumbent or climbing, green, 

conspicuously ringed with leaf scars, sometimes wholly or partially covered with fibrous leaf bases; unarmed. Leaves bifid or 

pinnate (our material), rarely entire, reduplicate; leaf sheaths 

closed or becoming split; leaf blades entire, bifid, and pinnately 

ribbed or regularly or irregularly pinnately divided (our material); leaflets (pinnae) few or many. Inflorescences spicate or 

1(2)-branched (our material), inter- or infrafoliar, 1 to several 

per leaf axil. Flowers unisexual, 3-merous. Staminate flowers: 

sepals 3, united basally or free; petals 3, free or variously connate; stamens 6, sometimes with staminodes; filaments short. Pistillate flowers: sepals 3, united basally or free; petals 3, usually 

connate; staminodes present and toothlike or absent; gynoecium 

3-locular, 3-ovulate; stigmas 3, recurved. Fruits 1-seeded, globose or oblong, black at maturity. Seeds globose or ellipsoid; 

endosperm homogeneous; embryo basal or lateral. [Arecoideae: 

Chamaedoreeae.]

A neotropical genus of 80–110 species found in Mexico, 

Central America, and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, and Brazil). Two species are known from 

Venezuela, both found in Guaramacal.

References. Cuenca and Asmussen-Lange (2007); 

Cuenca et al. (2008, 2009); Hodel (1992); Thomas et al. (2006).

Chamaedorea is the most species-rich genus of palm in the 

Americas. Molecular data (Thomas et al., 2006; Cuenca and 

Asmussen-Lange, 2007) resolve the genus as monophyletic but 

fail to support the eight subgenera that Hodel (1992) recognized 

on floral characters.

KEY TO THE SPECIES OF CHAMAEDOREA

1a. Robust plants, stems 2–8 m × 2.5–5 cm; inflorescences infrafoliar, staminate inflorescences ≥3 per node; leaf blades 2–3.5 

m long; middle leaflets (pinnae) 30–65(–85) cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. linearis

1b. Delicate plants, stems 1–3 m × 0.5–2.5 cm; inflorescences infra- or interfoliar, staminate and pistillate inflorescences 1 per 

node; leaf blades 0.8–1.5 m long; middle leaflets (pinnae) 20–30 cm long . . . . . . . . . . . . . . . . . . . . . . . C. pinnatifrons

Chamaedorea linearis (Ruiz & Pav.) Mart., Hist. Nat. Palm. 2: 

5. 1823; Llamozas et al., Libro Rojo Fl. Venez. 328. 2003. 

Martinezia linearis Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 

297. 1798.

Robust, arborescent plants; stems (trunks) solitary, 2–8 m × 

2.5–5 cm. Leaves 3–8, pinnate; leaf sheaths closed, to 1.2 m 

long; leaf blades 2–3.5 m long; leaflets (pinnae) 25–65 per side, 

linear, lanceolate, generally curved, acuminate, middle leaflets 

30–65(–85) × 2–6.5(–12) cm. Inflorescences 1-branched, infrafoliar, erect; staminate inflorescences ≥3 per node; pistillate inflorescences 1 per node; rachillae whitish (in flower), green or 

yellowish-brown (in fruit). Fruits globose, 1–2 cm in diameter, 

red (black at maturity?).

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). In Venezuela, evidently restricted to the Andes 

(Lara, Mérida, and Trujillo) and the Sierra de Perijá (Zulia; Map 

24). In the park, found on the south slope of Guaramacal in 

Qda. Honda; 1,850–1,950 m.

Chamaedorea pinnatifrons (Jacq.) Oerst., Vidensk. Meddel. 

Dansk. Naturhist. Foren. Kjøbenhavn 1858: 14. 1859. Borassus pinnatifrons Jacq., Pl. Rar. Hort. Schoenbr. 2: 65, tt. 

247, 248. 1797.

Delicate, shrubby plants; stems (trunks) solitary, 1–3 m × 

0.5–2.5 cm. Leaves 3–10, pinnate; leaf sheaths closed 4

⁄5 their 

length, 6–35 cm long; leaf blades 0.8–1.5 m long; leaflets (pinnae) 7–15 per side, sigmoid or rarely almost lanceolate, apical 

segments bifid, middle leaflets 25–30 × 2–15 cm. Inflorescences 

1-branched, inter- or infrafoliar, staminate and pistillate inflorescences 1 per node, erect to curved (pendulous in fruit); rachillae reddish, orange or yellowish. Fruits globose or ellipsoid, 

0.4–0.6(–1) cm in diameter, orange or bright yellow, black at 

maturity.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Ecuador, Peru, Bolivia, and Brazil). Widespread in northern and western Venezuela (Anzoátegui, Aragua, Cojedes, Distrito Federal, Falcón, Lara, Mérida, Miranda, 

MAP 24. Chamaedorea linearis occurrence in Venezuela.
NUMBER 100 • 2 7

Portuguesa, Táchira, Trujillo, Yaracuy, and Zulia; Map 25). 

In the park, found on both slopes of Guaramacal; (1,550–)

1,800–2,350 m.

This is one of the most widespread and variable palm species in the Americas. Although it is found only at lower elevations in Guaramacal, Chamaedorea pinnatifrons otherwise is 

found at a greater range of elevations including ones exceeding 

those of the park.

Geonoma Willd.

Geonoma Willd., Sp. Pl. 4(1): 174, 593. 1805.

Monoecious, small to medium-sized plants; stems solitary 

or clustered, erect or decumbent; internodes often yellowish and 

smooth; unarmed. Leaves pinnate, divided or entire; leaf sheaths 

open, margins fibrous, glabrous or tomentose; leaf blades bifid 

or with 2 or 3 pairs of leaflets (pinnae) or irregularly divided. 

Inflorescences solitary, spicate or 1–3(–4)-branched, interfoliar 

or rarely infrafoliar; peduncles short to long, glabrous or tomentose. Flowers 3-merous, arising in depressions or holes on the 

peduncular rachis. Staminate flowers: sepals 3, distinct; petals 

3, connate for 2⁄3 their length, valvate; stamens (3) 6. Pistillate 

flowers: sepals 3, united basally and adnate to the receptacle; 

petals 3, connate, briefly adnate to the receptacle; staminodes 

6-toothed or 6-lobed; gynoecium 3-carpellate, but 2 carpels vestigial at anthesis, 1-locular, 1-ovulate; style tubular; stigmas 3. 

Fruits 1-seeded, ± globose, sometimes somewhat pointed, green, 

brown or purple-black. Seeds ± globose; endosperm homogeneous; embryo erect basal. [Arecoideae: Geonomateae.]

A neotropical genus of 68 species (140 taxa) found in Mexico, Central America, the West Indies, and South America (Colombia, Venezuela, Trinidad and Tobago, the Guianas, Ecuador, 

Peru, Bolivia, Brazil, and Paraguay). Approximately 20 species 

(25 taxa) are found in Venezuela.

References. Henderson (2011); Roncal et al. 

(2011); Wessels Boer (1968).

The three species of Geonoma found in Guaramacal are 

closely related and belong to a clade that includes five species 

found at higher elevations in Central America, the West Indies, 

and South America (Henderson, 2011).

KEY TO THE SPECIES OF GEONOMA

1a. Inflorescences unbranched, simple spikes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. lehmannii

1b. Inflorescences unbranched or branched (our material) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 

2a. Inflorescences 1-branched, interfoliar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. orbignyana

2b. Inflorescences 2-branched, infrafoliar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. undata

Geonoma lehmannii Dammer ex Burret, Bot. Jahrb. Syst. 63: 

180. 1930.

Only the nominate subspecies is found in Venezuela and 

Guaramacal.

Geonoma lehmannii subsp. lehmannii

Geonoma jussieuana auct., non Mart.; Dorr et al., Contr. U.S. Natl. Herb. 

40: 39. 2000 [2001].

Small plants, 1–2(–4) m tall; stems solitary or clustered. 

Leaves 5–15 per stem, undivided or irregularly pinnate, 30–80 

cm long; leaflets (pinnae) 3–5 per side, apical segments bifid. Inflorescences simple spikes, erect; peduncular bracts tubular, 9–50 

cm long; peduncles 12–88 cm long. Fruits 5–10 × 5–7 mm, olivegreen to black.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, known only from the Andes (Lara, Portuguesa, and Trujillo; Map 26). Common in the understory of 

montane and cloud forest throughout the park; (1,600–)1,800–

2,600 m.

Common Name. Virtud.

Venezuelan material of Geonoma lehmannii is geographically isolated and differs from the closest populations of G. 

lehmannii subsp. lehmannii in northern Colombia by a number 

of morphological characters. Nonetheless, Henderson (2011) 

chose not to give formal taxonomic recognition to the Venezuelan populations.

Specimens from Central America (Panama) are segregated as 

G. lehmannii subsp. corrugata A. J. Hend. and differ significantly 

MAP 25. Chamaedorea pinnatifrons occurrence in Venezuela.
2 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

from the South American ones in a number of morphological 

characters (Henderson, 2011), most notably longer peduncles.

Geonoma orbignyana Mart., Voy. Amér. Mér. 7(3): 22. 1847 

[1843]; ibid. 8, Atlas: t. 11, fig. 1. 1847 [1843] (“Orbigniana”), t. 22A. 1847 [1844] (“Orbigniana”); Morillo, in 

Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 390, 

foto 3. 2010.

Only the nominate subspecies is found in Guaramacal.

Geonoma orbignyana subsp. orbignyana

Geonoma jussieuana Mart., Voy. Amér. Mér. 7(3): 24. 1847 [1843]; ibid. 8, 

Atlas: t. 12 [as “15”], fig. 1. 1847 [1843], t. 23A. 1847 [1844].

Small plants, 1–3(–6) m tall; stems (trunks) solitary or clustered. Leaves 4–20 per stem, undivided or irregularly pinnate, 

5–80 cm long; leaflets (pinnae) 3–9 per side, apical segments 

bifid. Inflorescences 1-branched (our material) or rarely spicate, 

infrafoliar, erect; peduncular bracts tubular, 3–50 cm long; peduncles 30–80 cm long. Fruits 4–10 × 5–7 mm, slightly pointed 

apically, olive-green to black.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). In Venezuela, found in the Andes (Lara, 

Mérida, Portuguesa, Táchira, and Trujillo) and the Cordillera 

de la Costa (Aragua and Yaracuy; Map 27). In the park, found 

in the understory of montane and cloud forest on both slopes of 

Guaramacal; 1,800–2,600 m.

Following Henderson (2011), we now consider Geonoma 

jussieuana to be a synonym of G. orbignyana subsp. orbignyana; 

this is a reversal of earlier treatments of the genus (Wessels Boer, 

1968; Henderson et al., 1995; Dorr et al., 2000), where the 

former name was applied to what we treat here as G. lehmannii

subsp. lehmannii. Inflorescence branching, a character previously 

used to separate these taxa as species, evidently is inconsistent 

and has no geographical basis.

Henderson (2011) recorded a dozen morphological characters that serve to separate the nominate subspecies, which 

occurs in South America, from G. orbignyana subsp. hoffmanniana (H. Wendl. ex Spruce) A. J. Hend., which is restricted to 

Central America. In particular, Henderson (2011) distinguished 

G. orbignyana subsp. hoffmanniana from G. orbignyana subsp. 

orbignyana by its longer peduncles.

Geonoma undata Klotzsch, Linnaea 20: 452. 1847; Llamozas 

et al., Libro Rojo Fl. Venez. 337. 2003; Morillo, in Morillo 

et al., Bot. Ecol. Monocot. Páramos Venez. 2: 391, foto 4. 

2010, pro parte.

Only the nominate subspecies is found in the Andes of Venezuela and Guaramacal.

Geonoma undata subsp. undata

Geonoma weberbaueri Dammer ex Burret, Bot. Jahrb. Syst. 63: 221. 1930; 

Dorr et al., Contr. U.S. Natl. Herb. 40: 39. 2000 [2001]; Morillo, in 

Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 392, foto 5. 

2010.

Medium-sized plants, 2–8 m tall; stems (trunks) solitary. 

Leaves 4–17 per stem, undivided or irregularly pinnate, (17–)

60–100 cm long; leaflets (pinnae) 1–60 per side, apical segments 

bifid. Inflorescences (1–)2–3-branched, infrafoliar; peduncular 

bracts 7–40 cm long; peduncles 10–35 cm long. Fruits 4–15 × 

6–10 mm, black.

MAP 26. Geonoma lehmannii subsp. lehmannii occurrence in 

Venezuela.

MAP 27. Geonoma orbignyana subsp. orbignyana occurrence 

in Venezuela.
NUMBER 100 • 2 9

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). In Venezuela, found in the Andes (Barinas, 

Lara, Mérida, Portuguesa, Táchira, and Trujillo), the Sierra de 

Perijá (Zulia), and in the Cordillera de la Costa (Aragua, Carabobo, Distrito Federal, Falcón, Miranda, and Yaracuy; Map 28). 

In the park, found in cloud forest on the south slope and in subpáramo on the Fila de Agua Fría and below Páramo de Vicuyal; 

(1,600–)1,800–2,800 m.

Common Name. Moriche.

Geonoma undata s.l. as circumscribed by Henderson (2011) 

is found in Mexico, Central America, the West Indies, and South 

America. He recognized 10 subspecies after correlating geographical range with differences in vegetative morphology.

Prestoea Hook. f.

Prestoea Hook. f., in Bentham and Hooker f., Gen. Pl. 3: 875, 899. 1883, 

nom. cons.

Monoecious, small to moderate-sized arborescent plants; 

stems (trunks) clustered or rarely solitary, ringed with prominent or obscure leaf scars; trunk usually persisting at the base of 

old shoots; unarmed. Leaves pinnate (our species), rarely simple, 

4–12 per plant, usually displayed horizontally; leaf sheaths open, 

occasionally partially closed; leaflets (pinnae) few to many, variously shaped, generally opposite or loosely alternate, horizontally disposed. Inflorescences branched, rarely spicate, interfoliar, 

rarely infrafoliar, erect or arching. Flowers unisexual, 3-merous. 

Staminate flowers: sepals 3, free, briefly imbricate at base; petals 

3, free, valvate; stamens 6. Pistillate flowers: sepals 3, free, imbricate; petals 3, free, imbricate, briefly valvate apically; staminodes 

6; gynoecium 1-locular, 1-ovulate. Fruits globose or ellipsoid, 

purple to black at maturity. Seeds globose; endosperm ruminate, 

rarely homogeneous; embryo subbasal. [Arecoideae: Euterpeae.]

A neotropical genus of 11 species found in Central America, 

the West Indies, and South America (Colombia, Venezuela, Guyana, Ecuador, Peru, Bolivia, and Brazil). Five species occur in 

Venezuela.

References. Henderson (1999); Henderson and de 

Nevers (1988); Henderson and Galeano (1996).

Prestoea acuminata (Willd.) H. E. Moore, Gentes Herb. 9: 286. 

1963; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 393–394, fig. 1B, 2, foto 6. 2010. Oreodoxa 

acuminata Willd., Samml. Deutsch. Abh. Königl. Akad. 

Wiss. Berlin 1803: 252. 1806.

Only the nominate variety is found in Guaramacal.

Prestoea acuminata var. acuminata 

Llamozas et al., Libro Rojo Fl. Venez. 342. 2003.

Arborescent plants, to 15 m × 20 cm. Leaves compound; 

leaf sheaths partially closed, 45–60 cm long, reddish to purple, 

some almost black; leaf blades 2–2.6 m long; leaflets (pinnae) 

23–30 or more per side; central leaflets larger, 55–60 × 4–5.5 cm. 

Inflorescences to 1.3 m long, horizontal; peduncles 5–20 × 2–3 

cm, secondary branches to 50 cm long. Flowers white, 3–4.5 mm 

long when fruiting. Fruits 1–1.3 cm in diameter.

Found in Central America and South America (Colombia, 

Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, found in 

the Andes (Mérida, Táchira, and Trujillo) and the Cordillera de 

la Costa (Aragua, Distrito Federal, Falcón, Miranda, Monagas, 

Sucre, and Yaracuy; Map 29). In the park, found in cloud forest in 

Qda. Jirajara on the south slope of Guaramacal; 1,950–2,250 m.

MAP 28. Geonoma undata subsp. undata occurrence in Venezuela.

MAP 29. Prestoea acuminata var. acuminata occurrence in 

Venezuela.
3 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Two other varieties of this species are recognized, but neither one is found in Guaramacal. Prestoea acuminata var. dasystachys (Burret) A. J. Hend. & G. A. Galeano, also found in the 

Andes, is readily distinguished from the nominate variety by its 

perianth size in fruit (to 7 mm long) and its significantly larger 

(1.4–1.8 cm in diameter) ovoid or obovoid fruit. Prestoea acuminata var. montana (Graham) A. J. Hend. & G. A. Galeano, 

restricted to the West Indies, is distinguished from the other varieties by its ± terete (vs. angular) rachillae with superficial (vs. 

somewhat sunken) flowers and fruit.

Wettinia Poepp. ex Endl.

Wettinia Poepp. ex Endl., Gen. Pl. 243. 1837.

Catoblastus H. Wendl., Bonplandia (Hannover) 8: 104, 106. 1860.

Monoecious, shrubby to arborescent plants; stems (trunks) 

solitary or clustered, conspicuously ringed with leaf scars; unarmed; supported by a dense cone of stilt roots. Leaves pinnate, 

spreading; leaflets (pinnae) of 2 types, one type undivided, asymmetrical and narrowly elliptical in outline, apices toothed, and 

the other type divided to the base into narrow segments. Inflorescences spikes or racemes, infrafoliar, 3–8(–15) per node, the central bud larger, the lateral buds progressively smaller. Staminate 

flowers: sepals 3(4), briefly connate or free; petals 3(4), briefly 

valvate or free; stamens 6–20. Pistillate flowers: sepals 3(4), imbricate, free or connate basally; petals 3(4), imbricate, free or 

connate basally; gynoecium of 1(2) fertile and (1)2 abortive 

carpels; style basal; stigmas 3. Fruits 1(2)-seeded, prismatic or 

pyramidal, flattened apically. Seeds ellipsoid, globose, prismatic 

or pyramidal, albuminous; embryo basal and erect. [Arecoideae: 

Iriarteeae.]

Found in Central America (Panama) and South America 

(Colombia, Venezuela, Ecuador, Peru, Bolivia, and Brazil). The 

genus comprises 21 species, with only 1 species found north of 

the Orinoco River in Venezuela.

Reference. Moore and Dransfield (1978).

Wettinia praemorsa (Willd.) Wess. Boer, Pittieria 17: 185. 1988; 

Llamozas et al., Libro Rojo Fl. Venez. 349. 2003. Oreodoxa 

praemorsa Willd., Samml. Deutsch. Abh. Königl. Akad. 

Wiss. Berlin 1803: 253. 1806. Catoblastus praemorsus

(Willd.) H. Wendl., Bonplandia (Hannover) 8: 104. 1860.

FIGURE 24A–C

Arborescent plants, 5–15(–20) m tall; stems clustered, occasionally solitary. Leaves 4–6, ascending, grayish-green; leaf 

sheaths to 1.5 m long; leaf blades to 2 m long; leaflets (pinnae) 

alternate, undivided, 20–30 per side or occasionally divided into 

2–8 segments, glabrous, nerves prominent, ultimate leaf segments flabellate, toothed. Inflorescences pendent; spadix deeply 

cup-shaped or hooded, smooth, brown, coffee or sometimes purple. Infructescences pendulous. Fruits ~3.5 × 2.5 cm, ellipsoid, 

green, salmon or yellow when mature, pubescent.

Found in South America (Colombia and Venezuela). In Venezuela, found in the Andes (Lara, Mérida, Portuguesa, Táchira, 

and Trujillo), the Sierra de Perijá (Zulia), and the Cordillera de 

la Costa (Aragua, Carabobo, Cojedes, Distrito Federal, Falcón, 

Guárico, Miranda, and Yaracuy; Map 30). This palm occurs in 

cloud forest on the north slope of the park, where it is especially 

noticeable from Qda. Segovia to the Páramo de Guaramacal 

along the Boconó–Guaramacal road; 1,800–2,600 m.

Common Name. Mapora.

Bromeliaceae

L. J. DORR AND B. STERGIOS

Perennial herbs, acaulescent or caulescent, epiphytic, terrestrial or lithophytic, sometimes tank-forming. Roots present or 

not, when present modified as holdfasts in epiphytic and lithophytic species. Leaves simple, spirally arranged, polystichous 

or distichous, usually rosulate or distributed along a stem; leaf 

sheaths ± dilated below; leaf blades usually ligulate to narrowly 

triangular, margins entire, spinose or serrate, covered with peltate scales (at least when young); stipules wanting. Inflorescences 

terminal or rarely lateral, scapose or sessile, simple or compound, panicles, racemes, spikes, capitula or solitary pseudolateral flowers, usually bearing brightly colored bracts. Flowers 

bisexual (sometimes functionally unisexual), regular or nearly 

so, 3-merous, often showy; perianth heterochlamydeous. Sepals 

3, free or connate, aestivation convolute or imbricate. Petals 

3, free or connate, abaxial surface appendaged basally or not 

(naked). Stamens 6, in 2 series; filaments free or joined to the 

petals and/or to each other; anthers 2-thecate. Ovary superior to 

inferior, 3-carpellate, 3-locular; placentation axile; ovules anatropous, numerous. Style terminal, 3-parted; stigmas 3, usually 

MAP 30. Wettinia praemorsa occurrence in Venezuela.
NUMBER 100 • 3 1

spiral-conduplicate. Fruit septicidal or loculicidal capsules, berries, or multiple fleshy fruits (Ananas Mill.). Seeds winged, plumose or unappendaged, 1 to many per locule; embryo small, 

embedded in copious endosperm.

Found in subtropical and tropical North America (USA), 

Mexico, Central America, the West Indies, and South America 

(all country-level political units) and west tropical Africa (1 species); ~58 genera and ~3,200 species. In Venezuela, there are ~22 

genera and 374 species.

References. Barfuss et al. (2005); Givnish et al. (2007, 

2011); Holst (1994); Hornung and Gaviria (1999); Horres et 

al. (2000, 2007); Oliva-Esteva and Steyermark (1987); Smith 

(1957, 1971); Smith and Downs (1974, 1977, 1979); Smith and 

Till (1998); Smith and Wood (1975).

We follow Givnish et al. (2007), who recognize eight subfamilies of Bromeliaceae where traditionally there had been 

three. Representatives of four of these eight subfamilies are 

found in the park.

KEY TO THE GENERA OF BROMELIACEAE

1a. Margins of leaf blades entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Plants terrestrial (our species) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pitcairnia

2b. Plants epiphytic (our species) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Petal claws coherent and forming a tube that equals the sepals; flowers usually polystichous . . . . . . . . . . . . . 4

 4a. Petal claws with 2 basal scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mezobromelia

 4b. Petal claws without scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Guzmania

 3b. Petal claws free or with a very short tube exceeded by the sepals; flowers usually distichous . . . . . . . . . . . . . . 5

 5a. Petal claws with 2 basal scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vriesea

 5b. Petal claws without scales (our species) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 6a. Stamens longer than the petals in length, exserted; sepals symmetric, free or rarely connate . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tillandsia

 6b. Stamens shorter than or equal to the petals in length, not exserted; sepals asymmetric, free or nearly so

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Racinaea

1b. Margins of leaf blades serrate or armed with spines or prickles (sometimes these are very small and restricted to the base 

of the leaf blades) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

7a. Fruit a dry capsule; seeds winged; ovary superior or nearly superior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Puya

7b. Fruit a dry or ± fleshy berry; seeds without wings or appendages; ovary inferior . . . . . . . . . . . . . . . . . . . . . . . . . . 8

 8a. Epiphytic herbs (our species); inflorescences paniculate, conspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . Aechmea

 8b. Terrestrial herbs; inflorescences corymbose, nearly hidden at the base of lower leaves . . . . . . . . . . . . . . Greigia

Aechmea Ruiz & Pav.

Aechmea Ruiz & Pav., Fl. Peruv. Prodr. 47. 1794, nom. cons.

Acaulescent, epiphytic (our species) or terrestrial herbs. 

Leaves densely rosulate; leaf sheaths usually well developed; leaf 

blades usually ligulate, margins long-spiny to spinulose-serrate. 

Inflorescences simple or compound, lax or dense, not strobilate; 

scape often well developed. Floral bracts entire, in 2 or more 

rows. Flowers distichous or polystichous, sessile or pedicellate, 

bisexual (our species) or rarely unisexual. Sepals often strongly 

asymmetric, free or shortly connate. Petals regular, free, appendaged (our species) or not; appendages 2, ± adnate to the petals. 

Stamens included, free or the second series adnate to the petals, 

unappendaged. Ovary inferior. Fruit a dry berry. Seeds unappendaged. [Bromelioideae.]

A poorly circumscribed neotropical genus of ~240 species 

found in Mexico, Central America, the West Indies, and South 

America (all major political units with the exception of Chile); 

~30 species occur in Venezuela.

References. Aguirre-Santoro and Betancur (2008); 

Faria et al. (2004); Givnish et al. (2011); Horres et al. (2007); 

Sass and Specht (2010); Smith and Downs (1979).

Aechmea, as currently circumscribed (e.g., Smith, 1971; 

Smith and Downs, 1979), is a highly polyphyletic genus (Smith 

and Till, 1998; Sass and Specht, 2010; Givnish et al., 2011), and 

there appear to be as many as six different origins of the species 

now assigned to what was once thought to be the largest genus in 

the Bromelioideae. Clades reflect geographic distribution rather 

than morphological similarity among the taxa of Aechmea and 

closely related genera analyzed by Sass and Specht (2010). The 

phylogenetic relationships of the sole species of Aechmea found 

in Guaramacal, which has not been sampled in molecular studies, are unknown, but we suspect this species will eventually 

be included in one of the Central American or northern South 

American clades instead of the Brazilian clades identified by Sass 

and Specht (2010).

Aechmea spectabilis Brongn. ex Houllet, Rev. Hort. 47: 311, t. 

1875.

Aechmea paniculigera auct., non (Sw.) Griseb.; Ortega et al., BioLlania 5: 35. 

1987; Dorr et al., Contr. U.S. Natl. Herb. 40: 40. 2000 [2001].

Robust, tank-forming epiphytic herbs. Leaves densely rosulate; leaf sheaths elliptic, densely brown lepidote; leaf blades ligulate, 0.8–1(–1.4) m × 6–8(–11) cm, margins laxly serrate with 
3 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

spines to 3 mm long, apex acute with a thick terminal cusp. Inflorescences panicles, 0.6–1 m long, broadly pyramidal, lax, greatly 

branched; scape flocculose, soon glabrous; primary bracts erect, 

broadly lanceolate, acute, longer than the internodes. Spikes 

laxly 1–5-flowered. Floral bracts minute. Sepals 9–12 mm long, 

strongly asymmetric, shortly connate at base, pale rose. Petals 

~25 mm long, bright red to purple; crenate scales present at base. 

Ovary ~7 mm long, narrowly obovoid or clavate.

Found in South America (Colombia and Venezuela). In Venezuela, known from the Andes (Lara, Mérida, Táchira, and Trujillo) and the Sierra de Periá (Zulia; Map 31). In the park, found 

in forest at lower elevations on the south slope of Guaramacal; 

(1,200–)1,600 m.

Aechmea spectabilis is easily recognized within the park as 

it is the only epiphytic bromeliad with baccate fruit; all the other 

epiphytic taxa have capsular fruit.

Greigia Regel

Greigia Regel, Index Seminum [St. Petersburg (Petropolitanus)] 1864, Suppl.: 

13. 1864.

Terrestrial or rarely epiphytic herbs; caudex short. Leaves 

spiral, imbricate the length of the stem; leaf blades narrowly 

triangular or linear, margins partially or completely serrate or 

serrulate with minute to conspicuous spines. Inflorescences simple or compound, usually axillary, densely spicate, reduced and 

subglobose, relatively few-flowered. Scapes very short. Flowers 

bisexual, sessile or subsessile. Sepals free or connate, symmetric. 

Petals connate 1⁄2 their length, usually without appendages. Filaments adnate to the petals. Ovary inferior. Berry fleshy. Seeds 

unappendaged. [Bromelioideae.]

A genus of 20–30 species found in Mexico, Central America, and South America (Colombia, Venezuela, Ecuador, Peru, 

Bolivia, Chile, and Argentina). Five species occur in Venezuela.

Reference. Smith and Downs (1979).

Greigia alborosea (Griseb.) Mez, in Martius, Fl. Bras. 3(3): 247. 

1891; Smith, Fl. Venez. 12(1): 285–287, t. 19. 1971; Morillo 

and Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 2: 406–407, fig. 1, foto 1. 2010. Nidularium alboroseum Griseb., Nachr. Königl. Ges. Wiss. Georg-AugustsUniv. 1864: 12. 1864.

Subcaulescent, terrestrial herbs. Leaves rosulate; leaf 

sheaths subelliptic, covered toward base below with ferrugineous scales; leaf blades ensiform, 0.8–1(–1.3) m × 1.8–2(–4) cm, 

margins serrate toward base with spines to 3 mm long, entire 

in the middle, and sparingly spinose-serrate above the middle, 

apex long-attenuate, pungent, glabrate above, white lepidote 

below. Inflorescences corymbose, axillary, 10–20-flowered, 

nearly hidden at the base of lower leaves. Outer floral bracts 

lanceolate, equaling the sepals. Sepals lance-ovate, ~2 cm long, 

apex mucronate, sharply carinate. Petals ~3 cm long, blades 

suberect to spreading, minutely cucullate. Ovary narrowly 

ellipsoid.

Endemic to Venezuela, where it is found in the Andes 

(Mérida, Táchira, and Trujillo) and the Cordillera de la Costa 

(Anzoátegui, Aragua, Distrito Federal, Miranda, and Sucre; Map 

32). In the park, found in cloud forest on the north slope of 

Guaramacal; 2,350 m.

This species is easily distinguished from other bromeliads 

in the park by its axillary inflorescences hidden among serrate 

leaves.

MAP 32. Greigia alborosea occurrence in Venezuela.

MAP 31. Aechmea spectabilis occurrence in Venezuela.
NUMBER 100 • 3 3

Guzmania Ruiz & Pav.

Guzmania Ruiz & Pav., Fl. Peruv. 3: 37. 1802.

Acaulescent or rarely caulescent, epiphytic or rarely terrestrial, often tank-forming herbs. Leaves densely polystichous; leaf 

sheaths conspicuous, imbricate; leaf blades mostly ligulate, margins entire. Inflorescences mostly bipinnate or simple, branches 

polystichously flowered; scape usually conspicuous, sometimes 

absent. Flowers sessile, bisexual. Sepals free to almost completely connate, symmetric or almost symmetric. Petals unappendaged, claws distinct from the blades, closely agglutinated 

(not truly connate) in a tube. Stamens included; filaments ± adherent (not adnate) to the petals. Ovary superior; style elongate; 

ovules many. Fruit a septicidal capsule. Seeds with a basal coma. 

[Tillandsioideae.]

A genus of 125–175 species found in the USA (Florida), 

Mexico, Central America, the West Indies, and South America 

(Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, and 

Brazil). In Venezuela, 28 species have been reported.

Reference. Smith and Downs (1977).

Guzmania appears to form a monophyletic clade within the 

Tillandsioideae (Barfuss et al., 2005; Givnish et al., 2011).

KEY TO THE SPECIES OF GUZMANIA

1a. Sepals exserted, not completely covered by the floral or primary bracts; leaf blades 40–50(–85) cm long . . . . . G. mitis

1b. Sepals completely covered by the floral bracts and sometimes also by the primary bracts; leaf blades 25–30 cm long . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. squarrosa

Guzmania mitis L. B. Sm., Contr. Gray Herb. 98: 31, t. 6, figs. 

4–5. 1932; Smith, Fl. Venez. 12(1): 256–257. 1971; Morillo 

and Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 2: 410. 2010.

Acaulescent tank-forming epiphytes, 1–1.5 m tall. Leaf 

sheaths broadly ovate, ± brownish at base; leaf blades ligulate, 

40–50(–85) × 3–5 cm, apex acute, punctulate-lepidote above 

and below. Inflorescences bipinnate, erect, 10–25 cm long, glabrous; primary bracts strict (rigidly erect), ovate or lanceolate, 

densely imbricate, foliaceous, longer than internodes; spikes 

globose or thick-ovoid, short-stipitate, 2.5–3 cm long, densely 

12–15-flowered. Floral bracts broadly ovate, shorter than sepals, 

obtuse. Sepals obovate, ~1 cm long, shortly connate, obtuse. Petals elliptic, ~2 cm long, blades erect, white.

Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, and Ecuador). In Venezuela, found at higher elevations in the western states (Falcón, 

Lara, Mérida, Portuguesa, Táchira, Trujillo, and Zulia; Map 33). 

In the park, found in forest mostly on the north slope; 1,800–

2,350 m.

Guzmania squarrosa (Mez & Sodiro) L. B. Sm. & Pittendr., 

J. Wash. Acad. Sci. 43: 403. 1953 [1954]; Smith, Fl. Venez. 

12(1): 260. 1971; Holst, in Berry et al., Fl. Venez. Guayana 

3: 590, fig. 504. 1997; Morillo and Briceño, in Morillo et 

al., Bot. Ecol. Monocot. Páramos Venez. 2: 411–412, fig. 

3A, foto 3. 2010. Thecophyllum squarrosum Mez & Sodiro, Bull. Herb. Boissier, sér. 2, 4: 877. 1904.

MAP 33. Guzmania mitis occurrence in Venezuela. MAP 34. Guzmania squarrosa occurrence in Venezuela.
3 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Guzmania monostachia auct., non (L.) Rusby ex Mez.; Ortega et al., BioLlania 5: 35. 1987.

Caulescent, epiphytic or terrestrial herbs, 40–70(–100) cm 

tall. Leaves forming a large, almost flat rosette; leaf sheaths 

narrowly elliptic, covered with brown scales; leaf blades linear, 40–50(–70) × 5–7 cm, apex abruptly acute, subglabrous 

above, punctulate-lepidote below. Inflorescences bipinnate, ± 

erect, 20–40 cm long, glabrous; primary bracts ovate, densely 

imbricate, foliaceous with showy red to orange (rarely yellow) 

striations, spreading to decurved at apex; spikes short, densely 

4–6-flowered glomerules. Floral bracts elliptic, shorter than sepals, obtuse or truncate. Sepals broadly elliptic, ~2 cm long, connate most of their length. Petals obtuse, ~2 cm long, yellow.

Found in South America (Colombia, Venezuela, Guyana, 

Ecuador, Peru, and Bolivia). In Venezuela, found at higher elevations in the Andes (Lara, Mérida, Portuguesa, Táchira, and 

Trujillo), the Sierra de Perijá (Zulia), the Cordillera de la Costa 

(Aragua and Distrito Federal), and the Venezuelan Guayana 

(Amazonas and Bolívar; Map 34). Found in forest and subpáramo throughout the park; 1,850–3,100 m.

Mezobromelia L. B. Sm.

Mezobromelia L. B. Sm., Proc. Amer. Acad. Arts 70: 151. 1935.

Medium-sized to large, tank-forming, acaulescent herbs, epiphytic (our species) or rarely terrestrial. Leaves rosulate; sheaths 

large; blades entire. Inflorescences compound, inflorescence 

branches polystichously or rarely distichously flowered; scape 

usually well developed. Flowers sessile or short-pedicellate; bisexual. Sepals ± free or connate, symmetric. Petals tightly conglutinated to the middle, forming a tube below, appendaged; 

appendages 2 per petal, adaxial. Stamens included; filaments adherent to the petals. Ovary superior; stigma conduplicate-spiral. 

Fruit a capsule. [Tillandsioideae.]

A genus of 9 species found in Central America (Costa Rica 

and Panama), the West Indies, and South America (Colombia, 

Venezuela, Trinidad and Tobago, the Guianas, Ecuador, Peru, 

and Brazil). Two species occur in Venezuela.

References. Luther (1995); Rauh (1992); Smith 

and Downs (1977).

The genus Mezobromelia cannot be distinguished vegetatively from Guzmania, but in flower the former is distinguished 

from the latter by the presence of appendages at the adaxial base 

of its petals.

Mezobromelia capituligera (Griseb.) J. R. Grant, Phytologia 

74: 428. 1993; Holst, in Berry et al., Fl. Venez. Guayana 

3: 605, fig. 512. 1997. Tillandsia capituligera Griseb., Cat. 

Pl. Cub. 254. 1866. Vriesea capituligera (Griseb.) L. B. Sm. 

& Pittendr., J. Wash. Acad. Sci. 43: 402. 1953; Smith, Fl. 

Venez. 12(1): 240–241. 1971; Ortega et al., BioLlania 5: 

35. 1987.

FIGURE 24I

Vriesea capituligera var. lutea Steyerm., J. Bromeliad Soc. 32: 113. 1982.

Tillandsia compacta auct., non Griseb.; Ortega et al., BioLlania 5: 35. 1987.

Large, tank-forming epiphytes, 40–70(–250) cm tall. Leaf 

blades ligulate, 30–60(–80) × 4–7(–12) cm, shorter than the 

scape, apex attenuate, glabrous above, obscurely punctulatelepidote below. Inflorescences erect, 30–50(–80) cm tall; primary 

bracts spreading, red or scarlet, showy. Sepals and floral bracts 

orange or red. Petals ~3 cm long, yellow or yellow-orange.

Found in the West Indies and South America (Colombia, 

Venezuela, Trinidad and Tobago, Ecuador, Peru, and Bolivia). 

Widespread in Venezuela (Amazonas, Aragua, Bolívar, Distrito 

Federal, Falcón, Lara, Mérida, Miranda, Portuguesa, Táchira, 

Trujillo, Yaracuy, and Zulia; Map 35). Found in forest throughout the park; 1,500–2,600 m.

Pitcairnia L’Hér.

Pitcairnia L’Hér., Sert. Angl. 7. 1788 [1789], nom. cons.

Acaulescent to caulescent, mostly terrestrial (our species) 

or rarely lithophytic or epiphytic herbs. Leaves polystichous or 

rarely distichous, fascicled or in dense spirals along the stem; 

leaf sheaths small, sometimes bulbous-thickened; leaf blades linear to broadly lanceolate, entire (our species) or serrate, attenuate or appearing petiolate. Scape present, often well developed. 

Inflorescences simple or compound. Floral bracts conspicuous 

to minute. Flowers long-pedicellate to sessile, showy, bisexual. 

Sepals convolute, free, attenuate to obtuse. Petals free, long and 

narrow, usually zygomorphic, convergent over the stamens, unappendaged or with a single scale (or scale remnant) at base, 

red, rarely white or yellow, very rarely blue or violet. Stamens 

elongate, either slightly shorter or slightly longer than the petals. Ovary almost wholly superior to wholly inferior; style long 

and slender; ovules numerous. Fruit a capsule. Seeds numerous, 

bicaudate or winged, rarely unappendaged. [Pitcairnioideae.]

MAP 35. Mezobromelia capituligera occurrence in Venezuela.
NUMBER 100 • 3 5

An overwhelmingly neotropical genus of ~280 species found 

in Mexico, Central America, the West Indies, and South America 

(Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, Brazil, 

Paraguay, and Argentina); 1 species native to west tropical Africa. 

There are ~41 species in Venezuela, 24 of which are endemic.

References. Givnish et al. (2004); Hornung-Leoni 

and Gaviria (2013); Smith and Downs (1974).

The sole African species, Pitcairnia feliciana (A. Chev.) 

Harms & Midlbr., appears to have reached tropical West Africa 

fairly recently (12–9.3 mya) via long-distance dispersal (Givnish 

et al., 2004, 2007, 2011). It also is the sole representative of the 

family to occur naturally outside the Americas.

KEY TO THE SPECIES OF PITCAIRNIA

1a. Leaf blades sessile, sometimes abruptly attenuate but not appearing petiolate, >2 cm wide; inflorescences racemose, 

>10 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Floral bracts showy, broadly ovate, exceeding the pedicels in length . . . . . . . . . . . . . . . . . . . . . . . . . . . P. maidifolia

2b. Floral bracts reduced, stipuliform, at most equaling the pedicels in length . . . . . . . . . . . . . . . . . . . P. brevicalycina

1b. Leaf blades appearing petiolate, <2 cm wide; inflorescences strobilus-like, <10 cm long . . . . . . . . . . . . P. hitchcockiana

Pitcairnia brevicalycina Mez, Repert. Spec. Nov. Regni Veg. 

16: 9. 1919; Smith, Fl. Venez. 12(1): 75–76. 1971; Morillo 

and Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 2: 414, fig. 4A. 2010.

Caulescent, terrestrial herbs, 1.5–1.8 m tall; stems often 

prostrate. Leaves persistent; leaf sheaths ovate, tomentuloselepidote, becoming glabrous; leaf blades lanceolate, 1–1.5 m × 

2–5 cm, base sometimes abruptly attenuate, not appearing petiolate, minutely lepidote, appearing glabrous. Scape strict; scape 

bracts triangular, shorter than the internodes and leaving much 

of the scape naked. Inflorescences simple, dense many-flowered 

racemes, 0.5–1.5 m long. Floral bracts greatly reduced, equal to 

or shorter than the pedicels. Sepals oblong, ~1 cm long. Petals 

broadly ligulate, 3.5–4 cm long, whitish to greenish-yellow or 

pale yellow, unappendaged or with traces of a scale.

Found in South America (Venezuela, Ecuador, and Peru). 

In Venezuela, restricted to the higher elevations in the western 

states (Barinas, Falcón, Lara, Mérida, Portuguesa, Táchira, Trujillo, and Zulia; Map 36). Open areas in cloud forest and subpáramo throughout the park; (1,450–)1,600–2,800 m.

In Guaramacal, hummingbirds visit the flowers of this species (pers. obs.).

Smith and Downs (1974) alluded to the fact that Pitcairnia caulescens K. Koch ex Mez, described from garden material 

that may have originated in Venezuela, might be conspecific with 

P. brevicalycina. If this is true, the former name is the one that 

would have to be adopted for our species.

Pitcairnia hitchcockiana L. B. Sm., Phytologia 5: 44, t. 6, figs. 

1–3. 1954.

More or less acaulescent, terrestrial or rupicolous herbs, 

0.5–0.6 m tall; often growing in dense aggregations. Leaves persistent; leaf sheaths narrowly ovate, lepidote, becoming glabrous; 

leaf blades linear-lanceolate, ~0.8 m × 1.5–2 cm, appearing 

MAP 36. Pitcairnia brevicalycina occurrence in Venezuela. MAP 37. Pitcairnia hitchcockiana occurrence in Venezuela.
3 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

petiolate, floccose when young, becoming glabrous. Scape slender, shorter than the leaf blades; scape bracts erect, foliaceous, 

exceeding the internodes in length. Inflorescences simple, slenderly ellipsoid, strobilus-like, few-flowered, ~10 cm long. Floral 

bracts imbricate, much longer than pedicels, red or scarlet. Sepals lanceolate, ~2 cm long. Petals ~4 cm long, red with a narrow 

white margin, appendaged; scale (appendage) at base.

Restricted to South America (Venezuela and Ecuador). In 

Venezuela, found only in the Andes (Táchira and Trujillo; Map 

37). In the park, found in forest on sandstone formations in 

sector El Santuario and near La Cueva below Cerro El Diablo; 

1,800–2,000 m.

Pitcairnia maidifolia (C. Morren) Decne. ex Planch., Fl. Serres 

Jard. Eur. 9: 151. 1854; Smith, Fl. Venez. 12(1): 63–65. 

1971; Holst, in Berry et al., Fl. Venez. Guayana 3: 644, fig. 

556. 1997; Morillo and Briceño, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 2: 414, fig. 4C, foto 4. 2010. 

Puya maidifolia C. Morren, Ann. Soc. Roy. Agric. Gand 5: 

453, t. 289. 1849.

Terrestrial or rupicolous herbs, to 1.5 m tall. Leaves rosulate, fasciculate, persistent; leaf sheaths narrowly ovate, floccose, 

becoming glabrous; leaf blades lanceolate, 0.5–1 m × 5–6 cm, 

appearing petiolate, glabrous. Scape erect; lower scape-bracts 

foliaceous, exceeding the internodes, upper scape-bracts ovate, 

acuminate. Inflorescences simple, dense (preanthesis) to lax 

(postanthesis) racemes, few- to many-flowered, 10–45 cm long. 

Floral bracts broadly ovate, shorter than the sepals, green or yellow and often tinged with red. Sepals broadly elliptic, asymmetric, 2.5–3 cm long. Petals linear, unequal, 5–6 cm long, white or 

greenish-white, unappendaged.

Found in Central America and South America (Colombia, 

Guyana, Suriname, Ecuador, and Brazil). Widespread in Venezuela (Amazonas, Aragua, Barinas, Bolívar, Distrito Federal, 

Lara, Mérida, Portuguesa, Táchira, Trujillo, and Yaracuy; Map 

38). In the park, found at lower elevations on the south slope of 

Guaramacal; 1,500–1,700 m.

Puya Molina

Puya Molina, Sag. Stor. Nat. Chili 160, 351. 1781 [1782].

Acaulescent or more commonly caulescent, terrestrial or 

lithophytic herbs; stems unbranched or branched. Leaves rosulate to densely polystichous, coriaceous; leaf sheaths distinct; leaf 

blades narrowly triangular, margins armed with spines or serrations, base never constricted. Scape usually present and well developed. Scape bracts foliaceous, densely imbricate, caducous or 

persistent. Inflorescences simple or compound, erect or nutant, 

sometimes massive (e.g., P. raimondii Harms). Flowers showy, 

bisexual. Sepals convolute, free, much shorter than the petals. 

Petals free, generally spreading at anthesis, spirally twisted after 

anthesis, unappendaged or with a pair of vertical folds (vestigial 

appendages). Stamens shorter than petals; filaments free, elongate. Ovary superior or nearly superior, glabrous. Style long and 

slender. Capsules loculicidal and usually also tardily septicidal. 

Seeds winged dorsally. [Puyoideae.]

A neotropical genus of ~200 species found in Central America (Costa Rica) and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Chile, and Argentina). Eight species 

occur in Venezuela.

References. Hornung-Leoni and Sosa (2006, 

2008); Jabaily and Sytsma (2013); Smith and Downs (1974).

Smith and Downs (1974) recognized two subgenera, Puya

subgen. Puya and P. subgen. Puyopsis (Baker) L. B. Sm.; the 

latter is distinguished from the former by having inflorescences 

and branches fertile throughout (vs. sterile toward the apex). 

This simple morphological classifycation is unlikely to be sustained as Jabaily and Sytsma (2013) using molecular characters found that Puya resolves into four major clades, including 

a central and northern Andes clade. Their data suggest that 

Puya originated in central Chile and moved north and that the 

northern Andes were colonized only once with no subsequent 

transitions back to the south. Overall, Puya is a classic example 

of a recent, rapid species-level radiation in the Andes (Jabaily 

and Sytsma, 2013).

KEY TO THE SPECIES OF PUYA

1a. Plants ≥1.5 m tall; inflorescences 2-branched, erect; sepals 2.5–3 cm long; petals white or whitish-green . . . . . . P. aristeguietae

1b. Plants <1 m tall; inflorescences simple, nutant (becoming erect in fruit); sepals 1.5–2 cm long; petals blue or greenish-blue

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. venezuelana

MAP 38. Pitcairnia maidifolia occurrence in Venezuela.
NUMBER 100 • 3 7

Puya aristeguietae L. B. Sm., Phytologia 7: 2, t. 1, figs. 9–12. 

1959; Smith, Fl. Venez. 12(1): 12–13. 1971; Morillo and 

Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 2: 418, fig. 5, foto 7. 2010.

FIGURE 24D, E

Puya venezuelana auct., non L. B. Sm.; Ortega et al., BioLlania 5: fig. 6. 1987.

Herbs to 3 m tall. Leaves rosulate, in dense clumps; leaf 

blades ensiform, 35–70 × 1.5–2.5 cm, rigidly coriaceous, margins armed with stout, dark spines. Scape erect, 1.5–3 m long; 

scape bracts broadly ovate, to 13 × 5 cm, long caudate, deciduous. Inflorescences 2-branched, erect. Sepals 2.5–3 cm long. Petals 5–6.5 cm long, white or yellow-green.

Found in South America (Colombia and Venezuela). In Venezuela, found in the Andes (Lara, Mérida, Táchira, and Trujillo) 

and in the Sierra de Perijá (Zulia; Map 39). Known from subpáramo and páramo in the park; (2,000–)2,700–3,100 m.

Common Name. Piñuela.

This species is a conspicuous element of the Páramo de 

Guaramacal and has also been found to the northeast of that 

locality on the Fila de Los Recostaderos. The type was collected 

southwest of Guaramacal in the Páramo de Guirigay.

Hummingbirds visit the flowers of Puya aristeguietae in the 

Páramo de Guaramacal (pers. obs.). Also, we strongly suspect 

that the spectacled bear, or “oso frontino” (Tremarctos ornatus), 

is feeding on the young leaf buds and shoots of this species in the 

same páramo as we have often observed young plants disturbed 

in a fashion suggestive of their being eaten.

Puya venezuelana L. B. Sm., Phytologia 7: 3, t. 1, figs. 13, 14. 1959.

FIGURE 24H

Herbs to 0.6(–1) m tall. Leaves rosulate, in compact clumps; 

leaf blades linear, 20–25 × 0.5–1 cm, subcoriaceous, margins 

serrulate, spines inconspicuous. Scape erect, to 0.75 m tall; scape 

bracts foliaceous, to 4.5 × 1.5 cm, caudate, persistent. Inflorescences simple, nutant (becoming erect in fruit). Sepals 1.5–2 cm 

long. Petals ~3 cm long, blue or greenish-blue.

Found in South America (Colombia and Venezuela). In Venezuela, known only from the Cordillera de Mérida (Mérida and 

Trujillo; Map 40). Páramo del Pumar; 2,600 m.

Within the park, this species seems to be restricted to 

Páramo del Pumar even though we reported earlier (Dorr et al., 

2000) that it was found on the Fila de Agua Fría. The presence of 

Puya venezuelana on this latter ridge was based on several sterile 

collections (i.e., Stergios & Zambrano 17654, 17664, 17680) 

that we now believe belong to two different taxa, one evidently 

a Puya (not P. venezuelana) and the other possibly a Pitcairnia.

Our material differs from the type of Puya venezuelana in 

several details and may, on further investigation, prove to be a 

distinct species. The Páramo del Pumar collections have serrulate (vs. spiny) leaves, scape bracts barely overlapping and exposing (vs. densely imbricate and concealing) the scape, and a 

cylindrical (vs. globose or strobilate) inflorescence. Puya venezuelana was described from the Páramo de Guirigay southwest of 

Guaramacal, and although it was long thought to be a Venezuelan endemic, Hornung-Leoni and Sosa (2006) extend its range 

into northern Colombia.

Racinaea M. A. Spencer & L. B. Sm.

Racinaea M. A. Spencer & L. B. Sm., Phytologia 74: 152. 1993.

Tillandsia subgen. Pseudocatopsis Baker, Handb. Bromel. 157, 192. [Aug–

Oct] 1889 (“Pseudo-Catopsis”).

Tillandsia sect. Pseudocatopsis André, Bromel. Andr. 62, 66. [Sept–Dec] 

1889 (“Pseudo-Catopsis”).

MAP 39. Puya aristeguietae occurrence in Venezuela. MAP 40. Puya venezuelana occurrence in Venezuela.
3 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Caulescent to short-caulescent, often tank-forming, epiphytic 

(our material) or rarely terrestrial herbs. Leaves rosulate; leaf 

sheaths large, sometimes forming a pseudobulb; leaf blades ligulate 

to subfiliform, margins entire. Scape bracts lanceolate, appressedascending, equal to or barely exceeding the internodes. Inflorescences simple or compound (our species), branches distichously 

flowered, rarely secund-flowered. Flowers actinomorphic, sessile 

to short-pedicellate, usually white to yellow, bisexual. Floral bracts 

equal to or shorter than the sepals. Sepals asymmetrical and broadest apically, free or rarely short-connate at base, convolute. Petals 

free, unappendaged, ± spreading, shorter or longer than the sepals. 

Stamens included; filaments free, sometimes adnate to the petals. 

Ovary completely superior. Style subsessile to short and stout. 

Stigma papillose, simple-erect type. Capsules septicidal. Seeds appendaged basally, coma white, pappiform. [Tillandsioideae.]

A neotropical genus of 60–70 species found in Mexico, Central 

America, and South America (Colombia, Venezuela, the Guianas, 

Ecuador, Peru, Bolivia, and Brazil). Ten species occur in Venezuela.

References. Barfuss et al. (2005, 2011); Smith and 

Downs (1977); Spencer and Smith (1993).

Spencer and Smith (1993) erected the genus Racinaea to 

accommodate a group of morphologically distinct taxa treated 

previously as Tillandsia subgen. Pseudocatopsis Baker (Smith, 

1971; Smith and Downs, 1977). Molecular data (e.g., Barfuss et 

al., 2005) have since confirmed that Racinaea is monophyletic, 

but the overall relationship of Racinaea with Tillandsia and other 

tillandsioid genera remains unresolved taxonomically. Accepting 

Racinaea and other monophyletic lineages as genera makes the 

remainder of Tillandsia paraphyletic. Fortunately, there are suggestions (Barfuss et al., 2011) that a reclassification of Tillandsia

and close relatives is forthcoming.

Racinaea can be distinguished from Tillandsia and other 

tillandsioid genera by its small, inconspicuous, and distichous 

flowers, free or nearly free asymmetrical sepals broadest toward 

the apex, included stamens and pistils, and short and stout 

styles.

KEY TO THE SPECIES OF RACINAEA

1a. Leaf blades ligulate; scape often greatly exceeding the leaves; floral bracts equal to or longer than the sepals . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. spiculosa

1b. Leaf blades lanceolate to narrowly triangular-lanceolate; scape ± equaling the leaves; floral bracts usually shorter than the 

sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. tetrantha

Racinaea spiculosa (Griseb.) M. A. Spencer & L. B. Sm., Phytologia 74: 157. 1993; Holst, in Berry et al., Fl. Venez. Guayana 3: 652–653. 1997; Dorr et al., Contr. U.S. Natl. Herb. 

40: 41. 2000 [2001]. Tillandsia spiculosa Griseb., Nachr. 

Königl. Ges. Wiss. Georg-Augusts-Univ. 1864: 17. 1864; 

Smith, Fl. Venez. 12(1): 189–190. 1971.

Three varieties of Racinaea spiculosa currently are recognized, and all three occur in Venezuela. Guaramacal material can 

be referred to the nominate variety.

Racinaea spiculosa var. spiculosa 

Holst, in Berry et al., Fl. Venez. Guayana 3: 653. 1997.

Acaulescent, epiphytic or lithophytic herbs, to 80 cm tall. 

Leaves rosulate; leaf sheaths broadly ovate, dark brown; leaf 

blades ligulate, (12–)20–40 × (2–)3.5–4 cm, apex roundedapiculate or acute, obscurely punctulate-lepidote. Scape 12–25 

cm tall, erect, often greatly exceeding the leaves; scape bracts 

narrowly elliptic, rounded-apiculate or short-caudate, ± equaling the internodes or the upper bracts slightly shorter. Inflorescences 2- or 3-pinnate, lax; primary bracts narrowly ovate, 

much shorter than the axillary branches. Spikes linear, lateral branches spreading, densely 7–10(–24)-flowered, usually 

long-stipitate. Floral bracts broadly ovate, equaling or slightly 

exceeding the sepals, greenish-red. Sepals broadly elliptic, 

0.3–0.7 cm long. Petals orange or yellow. Capsules cylindric, 

1.5–2.5 cm long.

Found in Central America (Costa Rica and Panama), the 

Greater Antilles, and South America (Colombia, Venezuela, 

Trinidad and Tobago, the Guianas, Ecuador, Peru, Bolivia, and 

Brazil). In Venezuela, found in the Andes (Mérida, Portuguesa, 

Táchira, and Trujillo), the Cordillera de la Costa (Aragua, Carabobo, Distrito Federal, Falcón, Miranda, Nueva Esparta, Sucre, 

and Yaracuy), and the Venezuelan Guayana (Amazonas and 

Bolívar; Map 41). In the park, found in cloud forest on both 

slopes of Guaramacal; 1,900–2,300 m.

MAP 41. Racinaea spiculosa var. spiculosa occurrence in Venezuela.
NUMBER 100 • 3 9

Racinaea tetrantha (Ruiz & Pav.) M. A. Spencer & L. B. Sm., 

Phytologia 74: 158. 1993; Holst, in Berry et al., Fl. Venez. 

Guayana 3: 653. 1997; Morillo and Briceño, in Morillo et 

al., Bot. Ecol. Monocot. Páramos Venez. 2: 423, fig. 8, foto 

12. 2010. Tillandsia tetrantha Ruiz & Pav., Fl. Peruv. 3: 39, 

t. 265. 1802; Smith, Fl. Venez. 12(1): 193–195. 1971.

Acaulescent, epiphytic herbs, 20–50(–80) cm tall. Leaves 

rosulate; leaf sheaths broadly elliptic-ovate, dark brown; leaf 

blades narrowly lanceolate to narrowly triangular-lanceolate, 

10–30(–45) × (0.5–)1.3–8 cm, apex acute or filiform, grayishgreen, often purple spotted, densely appressed-lepidote. Scape 

20–30 cm long, usually curved, ± equaling the leaves; scape bracts 

elliptic or oblong-elliptic, caudate to long-caudate, usually the 

sheathing bases shorter than the internodes. Inflorescences usually 2-branched, erect or pendulous, axis usually geniculate; primary bracts ovate to ovate-elliptic. Spikes spreading or reflexed, 

often becoming secund, 2–8(–11)-flowered, stipitate or subsessile. Floral bracts broadly ovate, shorter than the sepals. Sepals 

obovate or oblong, 0.5–1 cm long. Petals slightly exserted, conglutinated, yellow. Capsules cylindric, 2.5–3 cm long.

Found in Central America (Costa Rica), the Greater Antilles, 

and South America (Colombia, Venezuela, Ecuador, and Peru).

Racinaea tetrantha is a very variable species morphologically, and as many as seven varieties are recognized, including 

four in Venezuela. Our material can be assigned to either of two 

varieties distinguished by the following key.

KEY TO THE VARIETIES OF RACINAEA TETRANTHA

1a. Leaf blades narrowly triangular-lanceolate; primary bracts ~1⁄2 as long as the spikes; spikes distinctly stipitate . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. tetrantha var. caribaea

1b. Leaf blades ligulate; primary bracts exceeding the spikes in length; spikes subsessile . . . . . . . . R. tetrantha var. miniata

Racinaea tetrantha var. caribaea (L. B. Sm.) M. A. Spencer & 

L. B. Sm., Phytologia 74: 158. 1993; Holst, in Berry et al., 

Fl. Venez. Guayana 3: 654. 1997; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 41. 2000 [2001]. Catopsis fendleri Baker, 

J. Bot. 25: 175. 1887. Tillandsia fendleri (Baker) Mez, in 

de Candolle, Monogr. Phan. 9: 741. 1896 (“Fendleri”), non 

Griseb., 1865. Tillandsia caribaea L. B. Sm., Proc. Amer. 

Acad. Arts 70: 155. [Aug] 1935; L. B. Sm., Contr. Gray 

Herb. 106: 155. [Sep] 1935; Smith, Fl. Venez. 12(1): 191–

193. 1971. Tillandsia tetrantha var. caribaea (L. B. Sm.) 

Gouda, Fl. Guianas, ser. A, Phanerog. 3: 65. 1987. 

Racinaea tetrantha var. tetrantha auct., non (Ruiz & Pav.) M. A. Spencer 

& L. B. Sm.; Dorr et al., Contr. U.S. Natl. Herb. 40: 41. 2000 [2001].

Leaf blades narrowly triangular-lanceolate, 10–30(–45) 

× 1.3–2(–4.5) cm. Inflorescences bipinnate, composed of 7–13 

spikes; primary bracts ~1⁄2 as long as the spikes, red. Spikes 1.5–

3.5(–5) cm long, 2–8(–11)-flowered, distinctly stipitate.

Found in Central America (Costa Rica), the Greater Antilles, 

and South America (Venezuela and possibly Guyana). In Venezuela, found in the Andes (Lara, Mérida, Portuguesa, Táchira, and 

Trujillo), the Cordillera de la Costa (Aragua, Distrito Federal, 

Miranda, and Yaracuy), and the Venezuelan Guayana (Amazonas and Bolívar; Map 42). Found in cloud forest throughout the 

park; 1,700–2,400 m.

The leaves of this variety superficially resemble those of 

Tillandsia myriantha Baker, especially with respect to size and 

shape. In flower the two taxa are easily separated; the former has 

an inflorescence with several spreading or reflexed lateral spikes, 

whereas the latter has a terminal, subglobose inflorescence.

Racinaea tetrantha var. miniata (André) M. A. Spencer & L. B. 

Sm., Phytologia 74: 159. 1993; Dorr et al., Contr. U.S. Natl. 

Herb. 40: 41. 2000 [2001]. Tillandsia aurantiaca var. miniata André, Énum. Bromél. 7. 1888. Tillandsia tetrantha 

var. miniata (André) L. B. Sm., Contr. Gray Herb. 89: 15. 

1930; Smith, Fl. Venez. 12(1): 195. 1971.

Racinaea tetrantha var. scarlatina auct., non (André) M. A. Spencer & L. B. 

Sm.; Dorr et al., Contr. U.S. Natl. Herb. 40: 41. 2000 [2001].

Racinaea tetrantha var. A; Dorr et al., Contr. U.S. Natl. Herb. 40: 41. 2000 

[2001].

Leaf blades lanceolate, 20–30 × 2.5–4 cm. Inflorescences bipinnate, composed of 6–8 spikes; primary bracts exceeding the 

spikes in length, bright red or rarely pale greenish-yellow. Spikes 

~2.5 cm long, 2–4(–8)-flowered, subsessile.

Found in South America (Colombia, Venezuela, and Ecuador). In Venezuela, known only from the Andes (Lara, Táchira, 

and Trujillo; Map 43). Cloud forest and subpáramo throughout 

the park; 1,800–3,100 m.

MAP 42. Racinaea tetrantha var. caribaea occurrence in Venezuela.
4 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

This variety bears a superficial resemblance to Tillandsia 

biflora Ruiz & Pav., which can be distinguished by the progressively reduced size of the primary bracts from the base to the 

apex of an inflorescence. The primary bracts of Racinaea tetrantha var. miniata are not noticeably reduced in size.

Tillandsia L.

Tillandsia L., Sp. Pl. 286. 1753.

Acaulescent to short-caulescent, often tank-forming, epiphytic or rarely terrestrial or lithophytic herbs. Leaves rosulate 

or less often borne along the stem, polystichous or rarely distichous; leaf sheaths distinct, sometimes forming a pseudobulb; 

leaf blades ligulate to narrowly triangular or linear, margins 

entire, peltate or lepidote scales present (usually radially symmetric). Scape usually present, solitary (our species) or several, 

well developed to reduced or lacking, erect or pendulous. Inflorescences simple or compound, usually distichously flowered 

spikes or rarely reduced to a single flower, sometimes appearing 

polystichously flowered. Flowers sessile to short-pedicellate, bisexual. Floral bracts equal to or shorter than the sepals. Sepals 

asymmetric or rarely symmetric, free to slightly connate. Petals free, usually naked (without appendages) or appendaged. 

Stamens free or adnate to the base of the petals, rarely basally 

connate. Ovary superior. Stigma usually conduplicate-spiral 

or simple erect type, rarely convolute-blade or unique coralliform type. Capsules septicidal. Seeds cylindrical or fusiform, 

basally appendaged; appendage straight, plumose, and white. 

[Tillandsioideae.]

A neotropical genus of 600–650 species found in North 

America (USA), Mexico, Central America, the West Indies, and 

South America (all country-level political units). Approximately 

45 species are found in Venezuela.

References. Barfuss et al. (2011); Grant (1993); 

Smith and Downs (1977).

There is strong molecular support for the monophyly of 

Tillandsioideae and its division into four tribes (Barfuss et al., 

2005), but the circumscription of genera in this subfamily remains unsettled. Traditional morphologically based taxonomy 

relied on characters (e.g., spiral vs. distichous arrangement of 

the flowers, various connations of the corolla tube, inclusion vs. 

exsertion of stamens and styles, and presence vs. absence of petal 

appendages) that are of questionable diagnostic value (Grant, 

1993; Barfuss et al., 2005). Although molecular data make it 

clear that Tillandsia as circumscribed by Smith (1971) and Smith 

and Downs (1977) is not monophyletic, sampling remains insufficient to satisfactorily reclassify the genera of the subfamily. 

There are, however, suggestions (Barfuss et al., 2011) that such a 

reclassification is forthcoming.

KEY TO THE SPECIES OF TILLANDSIA

1a. Several to many scapes arising from a rosette of leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. complanata

1b. One scape arising from a rosette of leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Leaf blades very narrowly triangular, apex acute or filiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. myriantha

2b. Leaf blades ligulate, apex not acute or filiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Lateral spikes ascending, densely clustered near scape apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. compacta

 3b. Lateral spikes perpendicular to scape axis or if ascending not densely clustered near scape apex . . . . . . . . . . 4

 4a. Large epiphytes, (0.5–)1–3 m tall; leaf blades ≥60 cm long, ≥8 cm wide . . . . . . . . . . . . . . . . . . . T. fendleri

 4b. Small to medium-sized epiphytes, <0.5 m tall; leaf blades <30 cm long, <2.5 cm wide . . . . . . . . T. biflora

Tillandsia biflora Ruiz & Pav., Fl. Peruv. 3: 41, t. 268, fig. 

b. 1802; Smith, Fl. Venez. 12(1): 166–167. 1971; Morillo 

and Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 2: 425, fotos 13, 14. 2010.

Acaulescent, epiphytic herbs, 30–35 cm tall. Leaves rosulate; leaf sheaths ovate; leaf blades ligulate, 15–28 × 2–2.5 cm, 

apex acute, obscurely punctulate-lepidote, especially near the 

base, green and mottled with maroon or pink. Scape (16–)23–24 

cm long, usually curved; scape bracts foliaceous, densely imbricate, concealing the scape. Inflorescences bipinnate or rarely 

reduced to a single polystichous spike; primary bracts broadly 

ovate, inflated. Spikes laxly 1–3-flowered, short-stipitate. Floral 

MAP 43. Racinaea tetrantha var. miniata occurrence in Venezuela.
NUMBER 100 • 4 1

bracts broadly ovate, obtuse or rarely narrowly acute, much 

shorter than the sepals. Sepals oblong, ~1–1.5 cm long. Petals 

ligulate, ~1.5 cm long, blue, lavender or purple. Capsules cylindric, 3–4 cm long.

Found in Central America and South America (Colombia, 

Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, found in 

the Andes (Lara, Mérida, Táchira, and Trujillo), the Sierra de 

Perijá (Zulia), and the Cordillera de la Costa (Anzoátegui, Aragua, Distrito Federal, and Sucre; Map 44). In the park, found in 

forest and cloud forest on the north slope near El Cafenol and 

the Páramo de Vicuyal; 1,800–2,750 m.

Tillandsia compacta Griseb., Nachr. Königl. Ges. Wiss. GeorgAugusts-Univ. 1864: 18. 1864; Smith, Fl. Venez. 12(1): 

164–165. 1971; Holst, in Berry et al., Fl. Venez. Guayana 3: 

661. 1997; Morillo and Briceño, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 2: 425–426, foto 16. 2010.

Tillandsia compacta var. intermedia L. B. Sm., Phytologia 7: 3. 1959; Smith, 

Fl. Venez. 12(1): 165–166. 1971; Dorr et al., Contr. U.S. Natl. Herb. 

40: 41. 2000 [2001].

Acaulescent, epiphytic or, less common, terrestrial herbs, 30–

50(–70) cm tall. Leaves forming a narrowly crateriform rosette; 

leaf sheaths broadly ovate; leaf blades ligulate, 30–40 × 3–6 cm, 

apex broadly rounded and apiculate, punctulate-lepidote. Scape 

35–70 cm long, curved; scape bracts broadly elliptic, acute, red 

apically, erect, imbricate, concealing the scape. Inflorescences bipinnate, pendent or erect; primary bracts broadly elliptic, equaling or exceeding the lower spikes. Spikes (5–)7–10-flowered, 

sessile. Floral bracts broadly ovate, acute, erect, imbricate, exceeding the sepals. Sepals lanceolate, 1.5–2 cm long. Petals ~2 

cm long, purple. Capsules prismatic, ~3 cm long.

Found in the Greater Antilles and South America (Colombia, Venezuela, Ecuador, and Bolivia). Widespread in Venezuela, 

including the Andes (Mérida, Táchira, and Trujillo), the Sierra de 

Perijá (Zulia), the Cordillera de la Costa (Aragua, Distrito Federal, Falcón, Miranda, and Sucre), and the Venezuelan Guayana 

(Amazonas and Bolívar; Map 45). In the park, found in cloud 

forest on both slopes of Guaramacal; 2,200–2,900 m.

Tillandsia complanata Benth., Bot. Voy. Sulph. 173. 1844 

[1846]; Smith, Fl. Venez. 12(1): 172–173. 1971; Holst, in 

Berry et al., Fl. Venez. Guayana 3: 661. 1997; Morillo and 

Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 2: 426, fig. 9, foto 15. 2010.

FIGURE 24G

Acaulescent, epiphytic herbs, 30–35(–40) cm tall. Leaves in 

a dense rosette; leaf sheaths elliptic or narrowly ovate, densely 

and minutely brown lepidote; leaf blades ligulate, 20–40 × (1.5–)

3–5 cm, exceeding the scapes in length, abruptly apiculate or 

subobtuse, obscurely punctulate-lepidote, becoming glabrous 

above. Scapes numerous, 20–40 cm long, arising from the leaf 

axils, ascending then decurved; scape bracts narrowly lanceolate, acute or attenuate, erect, usually imbricate. Inflorescences 

simple, densely 4–24-flowered, complanate (i.e., flattened). Floral bracts elliptic, obtuse, erect, imbricate, exceeding the sepals, 

bright red or pinkish-purple. Flowers subsessile. Sepals lanceolate, 1–1.5 cm long. Petals ligulate, ~2 cm long, rose-red, purple 

or blue. Capsules narrowly cylindric, ~4 cm long.

Found in Central America (Costa Rica and Panama), the 

Greater Antilles, and South America (Colombia, Venezuela, 

Trinidad and Tobago, Guyana, Ecuador, Peru, Bolivia, and Brazil). Widespread in Venezuela and known from the Andes (Lara, 

MAP 44. Tillandsia biflora occurrence in Venezuela. MAP 45. Tillandsia compacta occurrence in Venezuela.
4 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Mérida, Portuguesa, Táchira, and Trujillo), the Sierra de Perijá 

(Zulia), the Cordillera de la Costa (Aragua, Distrito Federal, 

Falcón, Miranda, Monagas, and Yaracuy), and the Venezuelan 

Guayana (Amazonas and Bolívar; Map 46). Our records are 

from cloud forest and subpáramo throughout the park; 1,800–

2,750 m.

Plants of this species are easily recognized in the park because they typically have many axillary inflorescences that are 

distinctly complanate distally.

Tillandsia fendleri Griseb., Nachr. Königl. Ges. Wiss. GeorgAugusts-Univ. 1864: 17. 1864; Smith, Fl. Venez. 12(1): 

154–156, t. 10. 1971; Holst, in Berry et al., Fl. Venez. Guayana 3: 662, fig. 564. 1997.

FIGURE 24F

Tillandsia rubra var. reducta L. B. Sm., Fieldiana, Bot. 28: 151. 1951. Tillandsia fendleri var. reducta (L. B. Sm.) L. B. Sm., Mem. New York Bot. 

Gard. 14(3): 48. 1967; Smith, Fl. Venez. 12(1): 157–158. 1971; Holst, 

in Berry et al., Fl. Venez. Guayana 3: 662. 1997.

Acaulescent, epiphytic herbs, (0.5–)1–3 m tall; tank-forming. 

Leaves in a dense crateriform rosette; leaf sheaths subelliptic, 

densely brown punctulate-lepidote; leaf blades ligulate, 60–100 

× 8–11 cm, apex triangular-acuminate, glabrous, sometimes 

glaucous at base. Scape 1–1.5(–2) m long, erect; scape bracts foliaceous, erect, imbricate, and obscuring the scape. Inflorescences 

bipinnate, rarely simple or 3-pinnate, erect and showy; primary 

bracts like scape bracts. Spikes 10–30 × 5 cm, lanceolate to linear, complanate, densely 6- to many-flowered, ± stipitate with reduced sterile bracts at base. Floral bracts obovate, ± equaling the 

sepals. Flowers short-stipitate. Sepals oblong-lanceolate, 2.5–4.5 

cm long. Petals blue apically.

Found in the Greater Antilles and South America (Colombia, Venezuela, Trinidad and Tobago, Guyana, Ecuador, Peru, 

Bolivia, and Brazil). In Venezuela, found in the Andes (Lara, 

Mérida, Portuguesa, Táchira, and Trujillo), the Sierra de Perijá 

(Zulia), the Cordillera de la Costa (Anzoátegui, Aragua, Carabobo, Cojedes, Distrito Federal, Falcón, Miranda, Monagas, 

Sucre, and Yaracuy), and the Venezuelan Guyana (Amazonas 

and Bolívar; Map 47). This species occurs in forest on the north 

slope of the park; 1,800–2,200 m.

This is the largest of the Tillandsia species found in the park. 

It is one of the food plants of the spectacled bear, or “oso frontino” (Tremarctos ornatus; Goldstein, 2004; pers. obs.).

Tillandsia myriantha Baker, J. Bot. 25: 242. 1887; Smith, Fl. 

Venez. 12(1): 167–168. 1971.

Stoloniferous, epiphytic herbs, 20–30 cm tall. Leaves in a 

spreading rosette; leaf sheaths broadly ovate, cinereous-lepidote; 

leaf blades very narrowly triangular, 20–30 × 0.5–0.7(–1.5) cm, 

filiform-attenuate. Scape ~15 cm long, erect or curved; scape 

bracts ovate, imbricate. Inflorescences densely bipinnate, subglobose, 4–7 cm long; primary bracts erect, like the scape bracts, 

densely cinereous-lepidote. Spikes 4–6-flowered, elliptic, 2.5 cm 

long, subsessile. Floral bracts lanceolate, obtuse, ~1 cm long, 

equaling or slightly shorter than the sepals. Sepals short-connate, 

ovate, 0.7–0.9 cm long. Petals elliptic, ~1.5 cm long, blue or violet.

Found in South America (Colombia and Venezuela). In 

Venezuela, found in the Andes (Lara, Mérida, and Trujillo) and 

the Cordillera de la Costa (Anzoátegui, Aragua, Distrito Federal, Miranda, Nueva Esparta, Sucre, and Yaracuy; Map 48). In 

the park, found in forest and cloud forest on the south slope of 

Guaramacal; ~2,000–2,700 m.

MAP 46. Tillandsia complanata occurrence in Venezuela. MAP 47. Tillandsia fendleri occurrence in Venezuela.
NUMBER 100 • 4 3

Vriesea Lindl.

Vriesea Lindl., Edwards’s Bot. Reg. 29: ad t. 10. 1843, nom. cons.

Acaulescent, tank-forming, epiphytic (our material), terrestrial or lithophytic herbs. Leaves rosulate; leaf sheaths large; 

leaf blades usually ligulate, margins entire, inconspicuously lepidote, scales symmetrical. Scape usually conspicuous. Inflorescences simple or compound, usually distichously flowered spikes 

or rarely 1 or more polystichously flowered spikes; primary 

bracts generally conspicuous. Flowers bisexual; mostly shortpedicellate. Sepals convolute, free or almost free, symmetrical or 

subsymmetrical. Petals free or connate in a tube much shorter 

than the sepals, appendaged (2 scales at the base), firm and 

erect, mostly yellow (often with green tips) or white. Stamens 

included or exserted. Ovary usually superior or nearly superior; 

ovules many, usually caudate; stigma usually convolute-blade 

type, rarely simple-erect type. Capsule septicidal. Seeds fusiform; 

coma basal, long, and straight. [Tillandsioideae.]

A neotropical genus of 250–300 species found in Mexico, 

Central America, the West Indies, and South America (Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, and Argentina). Approximately 30 species are found in 

Venezuela.

References. Gomes-da-Silva et al. (2012); Smith 

and Downs (1977).

Although Vriesea has been treated as a natural group, its 

circumscription has been problematic because the morphological 

characters that have been used to define it (e.g., the presence of 

petal appendages and a connate corolla) are not unique within 

the Tillandsioideae (Gomes-da-Silva et al., 2012). In addition, 

phylogenetic analyses (Barfuss et al., 2005; Givnish et al., 2011) 

based on limited taxon sampling suggest that Vriesea is polyphyletic in origin. Nonetheless, molecular data indicate that Vriesea, Alcantarea (E. Morren ex Mez) Harms, and Werauhia J. R. 

Grant form a monophyletic clade within the Tillandsioideae, 

and they share a number of morphological characters (e.g., a 

partly inferior ovary, septicidal capsule, petal appendages usually 

present, and stigma mainly of the convolute-blade and cupulate 

types, rarely of the simple-erect type).

Vriesea incurva (Griseb.) Read, Phytologia 16: 458. 1968; Smith, 

Fl. Venez. 12(1): 219–220. 1971; Morillo and Briceño, in 

Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 430–

431, fig. 10A, foto 21. 2010. Tillandsia incurva Griseb., 

Nachr. Königl. Ges. Wiss. Georg-Augusts-Univ. 1864: 15. 

1864.

FIGURE 24J

Epiphytic herbs, (15–)20–45 cm tall. Leaves in a subglobose 

rosette; leaf sheaths broadly ovate to suborbicular, brown lepidote with a dense covering of tiny scales; leaf blades triangular, 

15–19(–35) × (1–)2–3 cm, apex long-attenuate, cinereous. Scape 

slender; scape bracts obovate or elliptic, ± imbricate. Inflorescences pendent, 20–40 cm long, simple or ± digitate with 2–5 

spikes; primary bracts scarcely larger than floral bracts. Spikes 

10–24 cm long, strongly complanate. Floral bracts elliptic, 2.5–

3.5 cm long, erect or spreading, greenish-yellow, margins pink. 

Sepals elliptic, 1.5–2 cm long. Petals ligulate, ~3.5 cm long, yellow or olive-green.

Found in Central America (Costa Rica and Panama), the 

Greater Antilles, and South America (Colombia, Venezuela, Guyana, Ecuador, and Bolivia). In Venezuela, collected in the Andes 

(Lara, Mérida, Portuguesa, Táchira, and Trujillo), the Sierra de 

MAP 48. Tillandsia myriantha occurrence in Venezuela.

MAP 49. Vriesea incurva occurrence in Venezuela.
4 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Perijá (Zulia), the Cordillera de la Costa (Aragua, Distrito Federal, Falcón, and Miranda), and the Venezuelan Guayana (Amazonas and Bolívar; Map 49). Found in forest on both slopes of 

Guaramacal; 1,800–2,600 m.

Burmanniaceae

L. J. DORR AND S. MIGUEL NIÑO

Achlorophyllous and mycoheterotrophic or sometimes autotrophic (some Burmannia L.) herbs; generally rhizomatous, 

sometimes with small tubers. Stem almost always solitary but also 

branched, often terete, colorless or yellowish in mycoheterotrophic taxa. Leaves alternate, simple, sessile, entire, sometimes rosulate, reduced and colorless or yellowish in mycoheterotrophic 

taxa, green in autotrophic taxa; stipules wanting. Inflorescences 

terminal, lax to dense, bracteate cymes, usually bifurcate, 1- to 

many-flowered. Flowers perfect, actinomorphic, sympetalous, 

composed of a basal tubular part and 6 tepals in 2 whorls. Floral 

tube cylindric, 3-angled or 3-winged, sometimes throat with an ornamented annulus. Stamens 3, opposite the inner tepals, inserted 

in the floral tube; anthers 2-thecate, introrse; with or without filaments, filaments very short. Gynoecium 3-carpellate, syncarpous; 

ovary inferior, 3-locular with axile placentation, 1-locular with 

parietal placentation or 3 free placental columns in the center of 

the ovary; ovules numerous, anatropous, minute; septal nectaries 

often present. Style cylindrical to filiform, as long as or shorter 

than the floral tube, 3-branched, rarely undivided. Fruit a capsule, 

winged or not, longitudinally dehiscent by slits or valves, or indehiscent. Seeds numerous, small, fusiform to subglobose.

A family of 8 genera and ~95 species found in tropical and 

subtropical regions worldwide, including North America (USA), 

Mexico, Central America, the West Indies, and South America. 

In Venezuela, 7 genera and 24 species occur.

References. Caddick et al. (2002a, 2002b); Jonker 

(1938); Maas and Maas-van der Kamer (1988); Maas et al. 

(1986); Maas-van de Kamer (1998); Merckx et al. (2006, 2008, 

2009, 2010); Wood (1983).

Burmanniaceae and Dioscoreaceae are the two families 

found in Guaramacal that belong to the Dioscoreales. Circumscription of both families has been problematic, with respect to 

not only which genera are to be included in each family but also 

the limits of some of these genera. In particular, the phylogenetic schemes of Caddick et al. (2002a, 2002b) and Merckx et 

al. (2010) present different hypotheses regarding relationships 

within this order, and we have decided to adopt the most recent 

hypothesis. In brief, Merckx et al. (2010) recovered seven wellsupported clades within Dioscoreales. Six of these clades were 

found to be sister to Burmanniaceae. The clades were Taccaceae, 

Thismiaceae, Trichopodiaceae, Afrothismia (Engl.) Schltr., Dioscoreaceae, and Nartheciaceae. Trichopodaceae, Taccaceae, and 

Afrothismia were each strongly supported monophyletic groups 

that also comprise a clade with Thismiaceae (Thismia Griff., 

Haplothismia Airy Shaw, and Tiputinia P. E. Berry & C. Wood., 

but not Oxygyne Schltr., were sampled). Consequently, our 

concept of Burmanniaceae, unlike most recent neotropical floras, 

excludes the genus Thismia, which sometimes is also treated as 

Burmanniaceae tribe Thismieae.

The achlorophyllous genera and species of Burmanniaceae 

usually are described as being saprophytic, but they are more 

accurately described as mycoheterotrophic as they have endomycorrhizae (Leake, 1994). Merckx et al. (2010) note that in 

the Dioscoreales, 14 of 23 genera include achlorophyllous mycoheterotrophic species, and they estimate that the mycoheterotrophic habit evolved at least six times. Furthermore, their data 

suggest that several extant mycoheterotrophic lineages are relatively ancient and appear to have Late Cretaceous origins. The 

achlorophyllous nature of Burmanniaceae and relatives generally 

confounded earlier efforts based on morphology to resolve phylogenetic relationships.

Gymnosiphon Blume

Gymnosiphon Blume, Enum. Pl. Javae 1: 29. 1827.

Cymbocarpa Miers, Proc. Linn. Soc. Lond. 1: 61. 1840.

Mycoheterotrophic herbs with cylindrical or slightly tuberous rhizomes. Stems branched or not. Leaves scalelike, ovate 

or subulate. Inflorescences lax to capitate, 2-branched cincinni, 

each cincinnus 1–24-flowered, sometimes reduced to a single terminal flower. Flowers erect or recurved, pedicellate or sessile. External tepals 3-lobed, internal tepals minute, sometimes swollen, 

inserted in the floral tube below the insertion of the outer tepals; 

floral tube not winged. Anthers 3, sessile, connective without 

or almost without appendages. Ovary 1-locular, with 3 parietal 

placentas, the partitions generally with glands. Style 3-branched 

apically, each branch with a hippocrepiform stigma (as seen from 

above), with or without 2 apical, tortuous, filiform appendages. 

Capsules ellipsoid, globose or obovoid, crowned by persistent 

perianth base, dehiscent either by 3 loculicidal valves separating 

from apex to base, 6 septicidal valves separating from base to 

apex or irregularly by withering of the fruit wall. Seeds ellipsoid 

to ovoid or fusiform.

A pantropical genus of ~30 species, the majority of which 

are found in the neotropics. In the Americas, the genus is known 

from Mexico, Central America, the West Indies, and South 

America (Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, and Brazil). Ten species occur in Venezuela, most in the 

Amazonian region.

Merckx et al. (2008) argue that diversification of Gymnosiphon began during the Eocene and that species migrated from 

the New to the Old World via boreotropical migration routes 

(i.e., high-latitude land connections when present-day northern 

regions supported tropical forest).

Gymnosiphon suaveolens (H. Karst.) Urb., Symb. Antill. 3: 438. 

1903; Maas and Maas, in Berry et al., Fl. Venez. Guayana 

3: 686. 1997. Benitzia suaveolens H. Karst., Linnaea 28: 

420. 1856 [1857].

Small, inconspicuous, achlorophyllous herbs, 10–30 cm tall. 

Stem whitish to purplish, 1–3 mm in diameter, glabrous. Leaves 
NUMBER 100 • 4 5

ovate, 1.5–3(–4.5) × 1.2–2(–3.5) mm. Inflorescences 2-branched 

cincinni, 2–3.5 cm long, erect or slightly recurved. Flowers white, 

7–15 mm long; floral tube with dark lines; pedicels 2–5 mm long. 

Style orange, 4–5 mm long; stigmas with contorted appendages 

to 5.5 mm long. Capsules ellipsoid to obovoid, 3.5–5 × 3 mm, 

longitudinally dehiscent, 3 valves separating from base to apex, 

valves persistent.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Ecuador, and Peru). In Venezuela, found 

mostly in montane forest in Aragua, Bolívar, Distrito Federal, 

Lara, Mérida, Miranda, Portuguesa, Táchira, Trujillo, and Zulia 

(Map 50). In the park, found in various localities on the south 

slope of Guaramacal; 1,700–2,350 m.

Gymnosiphon suaveolens is the only species of Gymnosiphon found in higher elevations of the Venezuelan Andes, and 

it is the only species of achlorophylous monocotyledon collected 

within the boundaries of the park.

Cannaceae

L. J. DORR AND S. MIGUEL NIÑO

Perennial herbs; rhizomes horizontal, usually branched; 

aerial stems well developed, unbranched; mucilage ducts present 

in rhizomes and stems. Leaves distichous or spiral; sheaths open, 

ligule absent; leaf blades with a prominent midvein and pinnateparallel veinlets, glaucous or lanuginose. Inflorescences terminal, 

bracteate, simple or compound thyrses; bracts of main axis subtending flowers or cincinni. Flowers perfect, epigynous, asymmetric, sessile or shortly pedicellate, red, purple, orange, yellow 

or white, generally showy. Sepals 3, much shorter than petals, 

imbricate, free, ± equal, green or sometimes colored. Petals 3, 

connate basally, sometimes free, generally unequal. Fertile stamen 1, petaloid, with a single marginal theca (sometimes appearing 2-thecate); staminodes 1–4(–5), variable-number petaloid, 

sometimes absent; inner staminode (labellum) usually wider than 

the other staminodes. Gynoecium 3-carpellate; ovary inferior, 

3-locular, warty to tuberculate; placentation axile; ovules few to 

numerous, anatropous; septal nectaries present. Style 1, petaloid; 

stigma decurrent along an edge of the style, papillate. Stamen, 

staminodes, and style united basally, forming a tube. Fruit a capsule, generally loculicidal, sometimes indehiscent, almost always 

crowned by the persistent sepals; fruit wall warty to tuberculate. 

Seeds globose to ellipsoid, numerous, black or dark brown, with 

a lid or collar; aril absent.

The family consists of a single tropical and subtropical genus 

of 10 species and various ornamental forms and hybrids originating in the Americas but now widely distributed throughout the 

world because of their appeal as ornamental or edible plants. In 

the Americas, they are found in North America (USA), Mexico, 

Central America, the West Indies, and South America (Colombia, 

Venezuela, the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, 

Argentina, and Uruguay). Four species occur in Venezuela.

References. Kubitzki (1998b); Maas-van der Kamer 

and Maas (2008); Prince (2010); Rogers (1984); Tanaka (2001).

Canna L.

Canna L., Sp. Pl. 1. 1753.

The characters of the genus are the same as those of the 

family.

Reference. Segeren and Maas (1971).

The center of diversity for the genus is South America, and 

thus, it is likely Canna originated there. This implies that the sole 

North American species is the result of long-distance dispersal 

(Prince, 2010).

The majority of Venezuelan species of Canna are given the 

common name “capacho.”

Canna jaegeriana Urb., Repert. Spec. Nov. Regni Veg. 15: 102. 

1917, nom. cons.

FIGURE 25A

Canna leucocarpa Bouché, Linnaea 18: 493. 1844 [1845], nom. rej.

Canna paniculata auct., non Ruiz & Pav.; Dorr et al., Contr. U.S. Natl. Herb. 

40: 42. 2000 [2001].

Canna sp.; Ortega et al., BioLlania 5: 36. 1987.

Plants to 3(–5) m tall. Leaf blades narrowly elliptic to narrowly ovate, 35–100 × 15–40 cm, base gradually decurrent on 

the petiole, apex acute to shortly acuminate, glabrous above, glabrous or ± lanuginose below. Inflorescences simple or branched, 

cincinni 1- or 2-flowered; bracts mostly caducous. Flowers erect, 

orange, 4–7.5 cm long; floral tube distinctly curved; the free part 

of staminodes erect; staminodes (3) 4 or 5. Capsules ellipsoid, 

4.5–10 × 1.5–4 cm. Seeds narrowly ellipsoid, 4–7 × 2–4.5 mm, 

shiny, black.

Found in the Greater Antilles and South America (Colombia, Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, known 

MAP 50. Gymnosiphon suaveolens occurrence in Venezuela.
4 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

from the Andes (Mérida, Táchira, Trujillo) and the Sierra de Perijá 

(Zulia; Map 51). Collected on the south slope in Qda. Jirajara near 

the village of Guaramacal and in El Santuario; ~1,600–1,950 m.

The upper surface of the leaves of Canna jaegeriana often 

turns dark brown to black in herbarium specimens.

Maas-van der Kamer and Maas (2008) noted that this species is often cultivated in Andean countries for its edible rhizome, 

but we have no evidence that it is used as food in Guaramacal.

Commelinaceae

L. J. DORR AND S. MIGUEL NIÑO

Perennial or sometimes annual herbs; usually terrestrial; 

erect or prostrate, sometimes climbing. Stems simple or branched, 

sometimes rooting from the nodes. Roots thin and fibrous; rhizomes, bulbs, stolons, and tubercles rare. Leaves basal or cauline, 

alternate, spiral or distichous, sometimes rosulate; leaf blades simple, entire, parallel-veined, ± succulent, sometimes purple-colored, 

usually sessile; sheaths closed, generally differentiated, sometimes 

perforated by the peduncle. Inflorescences terminal, terminal and 

axillary, or basal, cymose, thyrsiform (sometimes reduced) or 

umbel-like, rarely flowers solitary; sometimes enclosed in leaflike 

naviculiform or spathaceous bracts. Flowers heterochlamydeous, 

actinomorphic or zygomorphic, bisexual or andromonoecious, 

rarely gynomonoecious; 3-merous. Sepals (2) 3, free or sometimes 

connate, usually subequal, occasionally petaloid. Petals (2) 3, free 

or connate, equal or markedly unequal, colored, deliquescent. Stamens 6, sometimes fewer by abortion, in 2 series; anthers basifixed 

or dorsifixed, dehiscence longitudinal or rarely poricidal; filaments 

ciliate, sometimes colored; staminodes 2–4, often showy. Ovary 

syncarpous, superior, (2) 3-locular, sessile or shortly stipitate; placentation axile; ovules 1(2)-seriate, orthotropous; style 1, simple, 

usually slender; stigma 1, simple or rarely 3-lobed, enlarged or 

not; septal nectaries absent. Fruit a capsule, 2- or 3-valved, loculicidally dehiscent, rarely indehiscent or fruit fleshy; sometimes 

brightly colored. Seeds 1 to several per locule, dry.

A tropical and subtropical family of ~42 genera and ~650 

species, with representative taxa in temperate regions. The greatest species diversity is found in Africa and Madagascar. In Venezuela, 10 genera and ~30 species are native or naturalized.

References. Aristeguieta (1965); Burns et al. 

(2011); Evans et al. (2000, 2003); Faden (1998); Faden and 

Hunt (1991); Tucker (1989); Wade et al. (2006).

Most genera and species of Commelinaceae are assigned to 

the Commelinoideae, which is widespread geographically. Two 

genera and a dozen species are assigned to the Cartonematoideae, which is restricted to Australasia and continental Africa. 

Cartonematoideae generally lack raphide canals and glandular 

microhairs, which are found in Commelinoideae. Also, Cartonematoideae have yellow flowers, whereas Commelinoideae usually have pink or blue to white (rarely yellow to orange) flowers.

KEY TO THE GENERA OF COMMELINACEAE

1a. Flowers subtended by conspicuous naviculiform or spathaceous bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Leaves 20–25(–35) × 4–5(–8) cm; peduncles 13–14 cm long, terminated by 2 naviculiform bracts; flowers white; 

fertile stamens 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tradescantia

2b. Leaves 9–12 × 1.5–4 cm; peduncles 2.5–3 cm long, arising from the base of 1 spathaceous bract; flowers blue to 

white; fertile stamens 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Commelina

1b. Flowers subtended by small, inconspicuous bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Stems 8–40 cm long, rooting from the nodes; inflorescences cymose; petals white, pink or rose (rarely blue) . . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyodina

3b. Stems 1–3(–5) m long, not rooting from the nodes; inflorescences racemose; petals blue or purple . . . . . Dichorisandra

Commelina L.

Commelina L., Sp. Pl. 40. 1753.

Phaeosphaerion Hassk., Flora 49: 212. 1866.

Commelinopsis Pichon, Notul. Syst. (Paris) 12: 227. 1946.

Perennial or annual herbs; erect or creeping. Stems simple 

or slightly branched, glabrous. Roots usually fibrous. Leaves 

usually distichous; leaf sheaths well differentiated; leaf blades 

broadly to narrowly lanceolate, sessile or petiolate. Inflorescences terminal and leaf-opposed, composed of 1 or 2 cymes 

MAP 51. Canna jaegeriana occurrence in Venezuela.
NUMBER 100 • 4 7

enclosed in naviculiform or spathaceous bracts, proximal cyme 

several-flowered, distal cyme vestigial or with 1 to several staminate flowers. Flowers bisexual and staminate; markedly zygomorphic, pedicellate; bracteoles inconspicuous or absent. Sepals 

3, unequal, free or 2 apical sepals connate. Petals (2) 3, unequal, 

free, 1 apical petal generally small and clawed, sometimes absent, 2 basal petals larger, usually clawed. Fertile stamens 3; filaments free, glabrous; staminodes 2 or 3. Ovary 2- or 3-locular, 

glabrous; ovules 1 or 2 per locule, 1-seriate. Capsules (1-) 2- or 

3-valved, 2- or 3-locular, loculicidally dehiscent, occasionally indehiscent. Seeds 1(2)-seriate, 1 or 2 per locule, hilum linear.

A cosmopolitan albeit mainly tropical genus of ~170 species, with a center of diversity in southern Asia. In the Americas, 

found in North America (USA), Mexico, Central America, the 

West Indies, and South America (all country-level political units 

except Chile). Nine species occur in Venezuela.

Reference. Faden and Hunt (1987).

Throughout Venezuela species of Commelina of medium to 

small size are known by the common names “suelda con suelda” 

and “cucaracha.”

Commelina obliqua Vahl, Enum. Pl. 2: 172. 1805 [1806].

Commelina robusta Kunth, Enum. Pl. 4: 52. 1843.

Terrestrial herbs to 0.6 m tall; generally erect, occasionally 

creeping, often forming colonies. Stems glabrous to sparsely tomentose, 3–4 mm in diameter. Leaf sheaths 2–4 cm long, pubescent with hairs at the apex and sometimes in a vertical line along 

the sheath margin; leaf blades lanceolate to oblong-lanceolate, 

9–12 × 1.5–4 cm, bases attenuate or oblique, apices acuminate, 

puberulous above and below, silica crystals above appearing as 

translucent to whitish dots. Inflorescences clustered at apex of 

stem (solitary in juvenile plants); peduncles 2.5–3 cm long, pubescent; bracts spathaceous, 1.5–3 × 1.5–2 cm, green, glabrous 

or almost glabrous. Flowers short-pedicellate. Upper sepal small, 

~2 mm long, lower sepals larger, 3–5 mm long. Upper petals 

7–9 mm long, blue to white, lower petals reduced. Capsules 

4.5–7 mm in diameter; 2-valved, 3-locular. Seeds 1 or 2 per locule, pitted, brown.

Found in Mexico, Central America, and South America 

(Venezuela, Trinidad and Tobago, Guyana, French Guiana, Ecuador, Bolivia, Brazil, Paraguay, Argentina, and Uruguay); also 

recorded as an ephemeral weed in Europe (Belgium). Widespread 

in Venezuela (Aragua, Bolívar, Carabobo, Cojedes, Distrito Federal, Falcón, Lara, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo, and Zulia; Map 52). In the park, 

found in swampy areas near the Laguna de Aguas Negras and on 

the south slope near the village of Guaramacal; 1,850–2,000 m.

Dichorisandra J. C. Mikan

Dichorisandra J. C. Mikan, Del. Fl. Faun. Bras. t. 3. 1820, nom. cons.

Perennial herbs; erect, climbing, clambering or trailing 

vines (lianas). Stems simple or branched. Roots usually with 

distal tubers. Leaves spiral or distichous; leaf sheaths conspicuous, usually pubescent; leaf blades generally oblong-lanceolate, 

acute to acuminate, cuneate, pubescent or not. Inflorescences 

terminal, axillary or basal, thyrsiform or reduced; without 

spathaceous bracts. Flowers bisexual, sometimes staminate by 

abortion, ± actinomorphic, pedicellate, bracteolate. Sepals 3, 

subequal, free, unequal, the outermost longest, hooded, persistent in fruit, greenish or petaloid. Petals 3, subequal, free, blue 

or purple, glabrous. Stamens 5 or 6, ± equal or unequal; filaments short, glabrous; anthers elongate, 2-thecate, dehiscing by 

terminal pores. Ovary sessile, 3-locular; ovules 3–5 per locule. 

Capsules 3-valved, fleshy, dehiscent. Seeds 2-seriate, black, arillate; aril reddish.

A neotropical genus of 35–50 species found in Mexico, Central America, the Lesser Antilles, and South America (Colombia, 

Venezuela, Trinidad and Tobago, the Guianas, Peru, Bolivia, Brazil, Paraguay, and Argentina). Three species occur in Venezuela.

Reference. Aona (2008).

Dichorisandra hexandra (Aubl.) Kuntze ex Hand.-Mazz., Denkschr. Kaiserl. Akad. Wiss., Wien Math.-Naturwiss. Kl. 79: 

203. 1908; Berry, in Berry et al., Fl. Venez. Guayana 4: 359, 

fig. 269. 1998. Commelina hexandra Aubl., Hist. Pl. Guiane 

1: 35, t. 12. 1775.

FIGURE 4C–G

Climbing, clambering or scandent vines. Stems 2–3(–5) m 

long, glabrous to minutely puberulent. Leaf sheaths conspicuous, 2–2.6 cm long, pilose apically and with a line of hairs along 

the margins (fused edges); leaf blades elliptic to ovate-elliptic, 

12–20 × (2.5–)4.5–5 cm, bases acute to rounded or ± oblique, 

apices long-acuminate, glabrous above and below; sessile or 

short-petiolate. Inflorescences terminal, paniculate, 2–6(–10) cm 

long, ± globose; peduncles 2–3 cm long, pubescent; floral bracts 

0.5–1(–2) × 0.2 cm. Flowers few to many, violet to purple. Sepals 

5–8(–12) mm long. Petals 7–17 mm long. Stamens 6, all fertile. 

MAP 52. Commelina obliqua occurrence in Venezuela.
4 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 4. Commelinaceae. A, B. Tradescantia zanonia (L.) Sw. A. Habit. B. Flower. C–G. Dichorisandra hexandra (Aubl.) Kuntze ex Hand.-

Mazz. C. Habit. D. Floral bract. E. Flower. F. Anthers (note terminal pores). G. Fruit. (A, Dorr et al. 8726; B, Faden 76/164; C–G, Dorr et al. 

8519.)
NUMBER 100 • 4 9

Capsules obovoid, ~7 mm in diameter, purple, almost black, glabrous. Seeds 3–5; aril orange.

Found in Mexico, Central America, the Lesser Antilles, and 

South America (Colombia, Venezuela, Trinidad and Tobago, 

the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, and Argentina). Widespread in Venezuela (Amazonas, Apure, Aragua, 

Barinas, Bolívar, Carabobo, Cojedes, Delta Amacuro, Distrito 

Federal, Falcón, Guárico, Mérida, Miranda, Nueva Esparta, 

Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, and Zulia; Map 

53). In the park, found in forest understory on the south slope of 

Guaramacal; (1,200–)1,500–1,600 m.

Phyodina Raf.

Phyodina Raf., Fl. Tellur. 2: 16. 1836 [1837].

Perennial or rarely annual herbs; decumbent. Roots fibrous, 

rarely tuberous. Leaves spiral or distichous; leaf sheaths inflated, 

sparingly to densely pubescent throughout; leaf blades almost 

always succulent, usually pubescent, sessile. Inflorescences terminal or terminal and axillary, composed of pairs of sessile cincinni; cincinni subtended by small, inconspicuous bracts <1 cm 

long. Flowers bisexual, rarely gynomonoecious, actinomorphic, 

pedicellate or ± sessile; bracts and bracteoles small, scalelike, 

ciliate. Sepals (2–)4, subequal, free, papery, hyaline. Petals (2) 3, 

equal, free, not clawed, white, pink or rose (rarely blue). Stamens 

(3–)6, all fertile, equal or subequal; filaments bearded. Ovary 

2- or 3-locular; ovules (1) 2 per locule; stigma capitate. Capsules 

globose, 2- or 3-valved. Seeds (1) 2 per locule, hilum punctiform.

A monotypic, neotropical genus found in Central America 

and South America (Colombia, Venezuela, Ecuador, Peru, and 

Bolivia).

References. Bergamo (2003); Hunt (1986b).

We recognize Phyodina here in a narrow sense, including only P. gracilis (Kunth) Raf. The genus often is considered 

a synonym of Callisia Loefl., but the latter genus is polyphyletic (Hunt, 1986b; Bergamo, 2003). Molecular data (Bergamo, 

2003) suggest that Phyodina is more closely related to Tripogandra Raf. than it is to Callisia s. str. Although there is still 

uncertainty as to the limits of these and related tradescantioid 

genera, Phyodina can be distinguished from Tripogandra by its 

actinomorphic (vs. zygomorphic) flowers and equal or subequal 

(vs. alternating short and long) stamens.

Phyodina gracilis (Kunth) Raf., Fl. Tellur. 2: 16. 1836 [1837]. 

Tradescantia gracilis Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 

1: 208, t. 672. 1815 [1816]; ibid. [qu.] 1: 261. 1815 [1816]. 

Aneilema gracile (Kunth) Steyerm., Fieldiana, Bot. 28: 152. 

1951 (“gracilis”), non C. B. Clarke, 1881. Callisia gracilis

(Kunth) D. R. Hunt, Kew Bull. 38: 131. 1983.

Decumbent herbs; caespitose, generally forming small colonies in open areas. Stems 8–40 × 0.1–0.2 cm, almost always rooting at the nodes. Leaf sheaths 0.5–1 cm long; leaf blades narrowly 

to broadly ovate, 1–2.5(–4.5) × 0.5–1.5(–2) cm, bases rounded 

to slightly cordate, margins ciliate, apices apiculate, sparingly 

pubescent above and below. Inflorescences terminal and axillary; 

peduncles 1–5 cm long. Flowers pedicellate; pedicels 2–5 mm 

long, minutely glandular. Sepals 2–3 mm long, ovate, purple, pubescent with multicelled simple and glandular hairs. Petals ~4 

mm long, broadly ovate, white, pink or rose (rarely blue). Stamens 6, subequal. Ovary glabrous. Capsules ~2 mm in diameter. 

Seeds small, reticulate-pitted.

Found in Central America (Panama) and South America 

(Colombia, Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, 

MAP 53. Dichorisandra hexandra occurrence in Venezuela.

MAP 54. Phyodina gracilis occurrence in Venezuela.
5 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

restricted to the Andes (Mérida, Táchira, and Trujillo) and the 

Cordillera de la Costa (Distrito Federal; Map 54). In the park, 

found in disturbed areas along the Boconó–Guaramacal road 

and in pastures between Pozo Verde and the Páramo del Pumar; 

1,900–2,550 m.

Tradescantia L.

Tradescantia L., Sp. Pl. 288. 1753.

Campelia Rich., Démonstr. Bot. 46. 1808.

Zebrina Schnizl., Bot. Zeitung (Berlin) 7: 870. 1849.

Rhoeo Hance, Ann. Bot. Syst. 3: 659. 1852.

Setcreasea K. Schum. & Syd., Just’s Bot. Jahresber. 27(1): 452. 1901.

Cymbispatha M. Pichon, Notul. Syst. (Paris) 12: 224. 1946.

Perennial or rarely annual herbs; erect or ascendent. Stems 

branched or simple; nodes pronounced. Roots fibrous, sometimes 

tuberous. Leaves spiral or distichous; leaf sheaths well developed, 

pubescent; leaf blades broadly lanceolate to linear, sessile or 

rarely pedicellate. Inflorescences terminal or terminal and axillary, composed of pairs of cincinni; cincinni sessile, subtended by 

spathaceous or foliaceous bracts; bracteoles persistent. Flowers 

bisexual, ± actinomorphic; pedicellate or almost sessile. Sepals 

3, subequal, distinct, persistent in fruit. Petals 3, equal, free or 

connate basally, not or rarely clawed, pink, blue, violet or white 

(our species). Stamens 6, equal or subequal, all fertile; filaments 

bearded or glabrous. Ovary sessile, 3-locular, glabrous; ovules (1) 

2 per locule, 1-seriate. Capsules 3-valved, 3-locular, dehiscent. 

Seeds (1) 2 per locule, blackish, hard, hilum oblong to linear.

A temperate and tropical American genus of ~70 species 

found in North America (USA), Mexico, Central America, the 

West Indies, and South America (Colombia, Venezuela, Guyana, 

Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina, and Uruguay); adventive in Africa and Asia. Three species are native or 

naturalized in Venezuela.

References. Hunt (1975, 1986a).

In Venezuela, in addition to the three native or naturalized 

species of Tradescantia, at least three additional species are known 

to be cultivated (Campbell in Hokche et al., 2008), but there is no 

evidence that these latter species have become naturalized.

Tradescantia zanonia (L.) Sw., Fl. Ind. Occid. 1: 604. 1797; 

Berry, in Berry et al., Fl. Venez. Guayana 4: 363, fig. 274. 

1998. Commelina zanonia L., Sp. Pl. 41. 1753. Campelia 

zanonia (L.) Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 210. 

1815 [1816]; ibid. [qu.] 1: 264. 1815 [1816].

FIGURE 4A, B

Herbs, to 2 m tall; generally erect. Stems often simple, 3–5 

mm in diameter, somewhat succulent, usually glabrous. Leaves 

clustered at the apex of the stems; leaf sheaths 2–2.5 cm long, 

inflated, sparingly to densely pubescent especially at apex; leaf 

blades elliptic to narrowly elliptic, 20–25(–35) × 4–5(–8) cm, 

bases decurrent, margins ciliate, apices long-acuminate, glabrous 

(or sometimes sparingly to densely pubescent) above and below. 

Inflorescences axillary, composed of 1 pair of cymes and 2 spathes; 

peduncles simple or branched, 13–14 cm long; emerging from leaf 

sheaths; spathes 2–3.5 × 1–2 cm, naviculiform, ± open, green, 

margins ciliate, otherwise glabrous. Flowers white. Sepals 3–5 mm 

long, purple (accrescent in fruit). Petals 6–10 mm long, white. Filaments bearded or not. Fruit (including accrescent calyx) globose, 

3–5(–7) mm in diameter, fleshy, purple to black, shiny.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Ecuador, Peru, Bolivia, and Brazil); introduced in the West Indies. Widespread at higher elevations in 

Venezuela (Amazonas, Anzoátegui, Aragua, Barinas, Bolívar, 

Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, 

Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, 

and Zulia; Map 55). In the park, always in the understory 

and principally in humid areas on both slopes of Guaramacal; 

(1,200–)1,500–2,600 m.

This species is unique in the family in that the pedicel and 

calyx become fleshy and purplish in fruit, converting the normal 

tradescantioid capsule into a fleshy “berry.” This presumably is 

an adaptation to bird dispersal.

Tradescantia zanonia occasionally is cultivated as an ornamental; one common cultivar is a selection chosen for its variegated leaves.

Costaceae

L. J. DORR AND B. STERGIOS

Perennial, terrestrial (our species) or rarely epiphytic herbs; 

rhizomatous; nonaromatic. Indument of unicellular or multicellular, uniseriate hairs, or glabrous. Leaves simple, entire, spirally 

arranged; sheaths closed, tubular, ligulate; ligules truncate or 

2-lobed. Inflorescences open or dense strobiliform or capitate 

spikes, terminal on a leafy stem (our species) or on a separate 

MAP 55. Tradescantia zanonia occurrence in Venezuela.
NUMBER 100 • 5 1

leafless, basal shoot or flowers solitary in leaf axils; bracts imbricate, conspicuous, spirally arranged, with a linear nectariferous 

callus just below the apex, with or without a foliaceous appendage, 

subtending 1 (our species) or 2 flowers; bracteoles 1, naviculiform 

(our species) or tubular at the base. Flowers epigynous, perfect, 

zygomorphic. Calyx tubular, ± deeply (2) 3-lobed, lobes equal or 

sometimes unequal. Corolla 3-lobed, the medial lobe larger than 

the lateral ones, lobes connate basally, forming a distinct tube. Stamen 1; anther 2-thecate; filament often ± petaloid; labellum (i.e., 

fusion of 5 staminodes) large, petaloid, adnate to corolla, equal to 

or longer than corolla lobes, ± 3–5-lobed; basal part of the stamen 

and labellum united into a papillate tube. Ovary (2–)3-locular, inferior, septal nectaries 2, near the apex; placentation axile; ovules 

numerous, anatropous; style 1, filiform, in a slot between the thecae; stigma 1, 2-lamellate, with a dorsal, 2-lobed appendage or cupuliform and unappendaged, margins ciliate. Fruit a (2–)3-locular 

capsule, 3-angled, crowned by the persistent calyx, loculicidally 

dehiscent or indehiscent. Seeds numerous, angular or ellipsoid, 

black or dark brown, arillate; aril white or yellow.

A family of 7 genera and 70–100 species widely distributed 

in tropical and subtropical regions of the world. In the Americas, found in Mexico, the West Indies, and South America (Colombia, Venezuela, Trinidad and Tobago, the Guianas, Ecuador, 

Peru, Bolivia, Brazil, Paraguay, and Argentina). Three genera 

and 12 native or naturalized species are found in Venezuela.

References. Larsen (1998); Specht (2006); Specht 

and Stevenson (2006); Specht et al. (2001).

Costus L.

Costus L., Sp. Pl. 2. 1753.

Robust herbs, usually >1 m tall. Leaves short-petiolate; ligules usually large and conspicuous. Inflorescences terminal on 

a leafy stem (our species) or on a separate, leafless basal shoot 

directly from the rhizome, strobiliform (rarely capitate) spikes; 

bracts coriaceous (rarely chartaceous), green, yellow, orange or 

red, terminating in a foliaceous appendage or unappendaged; 

bracteoles naviculiform (our species) or tubular. Calyx lobes ± 

equal, rarely exceeding the bracts. Corolla white, yellow, orange or red. Labellum large and spreading or small and tubular. 

Stigma 2-labiate (our species) or cupuliform. Ovary 3-locular. 

Capsules ellipsoid to globose, tardily dehiscent or indehiscent 

and seeds released by decay of the fruit wall. Seeds black or dark 

brown, ellipsoid; aril conspicuous, lacerate, white.

A pantropical genus of 40–70 species, the vast majority of 

which are found in the Americas in Mexico, Central America, 

the West Indies, and South America (Colombia, Venezuela, the 

Guianas, Ecuador, Peru, Bolivia, and Brazil). Nine species occur 

in Venezuela.

References. Maas (1972, 1977).

KEY TO THE SPECIES OF COSTUS

1a. Inflorescence bracts green to red, foliaceous appendages green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. guanaiensis

1b. Inflorescence bracts dark red, without appendages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. spiralis

Costus guanaiensis Rusby, Bull. Torrey Bot. Club 29: 694. 1902 

(“Guanaiense”).

Three varieties of this neotropical species are currently recognized; the following is the only one that has been collected in 

Guaramacal.

Costus guanaiensis var. macrostrobilus (K. Schum.) Maas, Fl. 

Neotrop. Monogr. 8: 52. 1972; Maas, Fl. Venez. 11(2): 

239–241. 1982; Maas and Maas, in Berry et al., Fl. Venez. 

Guayana 4: 428, fig. 341. 1998. Costus macrostrobilus 

K. Schum., in Urban, Symb. Antill. 4: 159. 1903.

Costus guanaiensis var. tarmicus auct., non (Loes.) Maas; Maas, Fl. Neotrop. 

Monogr. 8: 57. 1920.

Plants 1–4(–6) m tall. Leaf blades narrowly ovate to narrowly obovate, 28–42 × 7–10.5 cm, apex shortly acuminate, 

base cuneate, rounded or rarely cordate, strigose and scabrid 

to the touch above, densely villous-puberulent below; sheaths 

± glabrous to noticeably villous-puberulent. Inflorescences 5–10

(–30) × 4–6(–10) cm; bracts with foliaceous appendages, broadly 

ovate, green to red. Corolla white, reddish-white or yellowishwhite, 70–100 mm long. Capsule 10–20 mm long, densely sericeous to glabrous.

Found in Mexico, Central America, the West Indies (Puerto 

Rico), and South America (Colombia, Venezuela, Trinidad and 

MAP 56. Costus guanaiensis var. macrostrobilus occurrence in 

Venezuela.
5 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Tobago, Guyana, Ecuador, Peru, and Brazil). Widespread in Venezuela (Amazonas, Anzoátegui, Apure, Aragua, Barinas, Bolívar, 

Delta Amacuro, Distrito Federal, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, and Zulia; 

Map 56). In the park, found in cloud forest on the north slope of 

Guaramacal; ~2,600 m.

Common Names. Ciruelo and saladillo.

The type of this variety comes from Puerto Rico, but because all other West Indian material is cultivated, Maas (1972: 

54) speculated that this variety is not native to the West Indies.

Costus spiralis (Jacq.) Roscoe, Trans. Linn. Soc. London 8: 350. 

1807. Alpinia spiralis Jacq., Pl. Rar. Hort. Schoenbr. 1: 1, 

t. 1. 1797.

Two varieties of this heterogeneous South American species 

are currently recognized; the nominate variety is the only one 

that has been found in Guaramacal and in Venezuela.

Costus spiralis var. spiralis

Maas, Fl. Venez. 11(2): 252–256. 1982; Maas and Maas, in 

Berry et al., Fl. Venez. Guayana 4: 428, fig. 344. 1998.

Plants 1–3.5(–4) m tall. Leaf blades narrowly elliptic, 

10–40 × 5–15 cm, apex acuminate, base cuneate to rounded, 

glabrous above and below; sheaths glabrescent. Inflorescences 

4–10(–20) × 2–5(–8) cm; bracts without appendages, broadly 

ovate, dark red. Corolla pinkish-red to salmon-red, 45–60 mm 

long. Capsule 10–15 mm long, glabrous.

Found in South America (Colombia, Venezuela, the Guianas, Peru, Bolivia, and Brazil); cultivated in the Greater Antilles 

(Cuba). Venezuelan records are from Amazonas, Apure, Aragua, 

Barinas, Bolívar, Falcón, Mérida, Miranda, Portuguesa, Táchira, 

Trujillo, Yaracuy, and Zulia (Map 57). Collected once on the 

south slope of Guaramacal in what appears to be the upper elevational limit of the variety; 1,500 m.

Costus spiralis var. villosus Maas, which is distinguished 

from the nominate variety by having leaves barely to densely villous, apparently is restricted to the Guianas and Brazil.

Cyclanthaceae

L. J. DORR AND B. STERGIOS

Perennial, monoecious, terrestrial, hemiepiphytic or epiphytic herbs or lianas. Caulescent or ± acaulescent; stems branched 

or unbranched, rhizomatous to aerial, usually ± woody. Leaves 

spirally arranged or distichous; mature leaves with open sheaths; 

petiolate (our species) or not; leaf blades usually bifid, rarely entire or divided into 4 segments, 1- or 3-costate. Inflorescences axillary or terminal, unbranched, pedunculate spadices subtended 

by 2–11 persistent or caducous, conspicuous, foliose spathes. 

Flowers unisexual, densely packed, generally (Carludovicoideae) 

in spirally arranged groups, each group consisting of 1 pistillate 

flower surrounded by 4 staminate flowers, or (Cyclanthoideae) in 

alternate cycles of staminate and pistillate units. Staminate flowers (Carludovicoideae): perianth lobes located on the periphery 

of the receptacle or only on the outer (abaxial) side, lobes usually 

in 1 whorl. Staminate flowers (Cyclanthoideae): linear rows of 

stamens; stamens numerous; anthers basifixed, 2-thecate, usually 

longitudinally dehiscent; filaments subconnate at the base, swollen to form basal “bulbs.” Pistillate flowers (Carludovicoideae): 

free or partially connate; tepals 4, free (our species) or connate, 

epigynous to perigynous, heteromorphic, subfleshy; staminodes 

4, filiform or subligulate; ovary 1-locular, 4-carpellate, exserted 

or partially buried in the rachis, with 4 parietal, subapical or 

apical placentae, or with 1 apical placenta, each placenta with 

numerous anatropous ovules; styles lacking or 4, free or ± concrescent and forming a common style; stigmas 4, alternating with 

the tepals; tepals, styles, and stigmas persistent. Pistillate flowers 

(Cyclanthoideae): not perceptible, coalescent into cycles. Fruits 

(Carludovicoideae): ± fleshy, free or united into a syncarp. Fruits 

(Cyclanthoideae): ± dry, united to form hollow rings filled with 

seeds. Seeds numerous, small to large, flat to terete.

A neotropical family of 12 genera and 200–230 species 

found in Mexico, Central America, the West Indies, and South 

America (Colombia, Venezuela, Trinidad and Tobago, the Guianas, Ecuador, Peru, Bolivia, and Brazil). Nine genera and ~40 

species occur in Venezuela.

References. Eriksson (1994); Harling (1958); Harling et al. (1998); Rudall and Bateman (2006).

Division of Cyclanthaceae into two subfamilies is based 

on a morphological phylogenetic analysis (Eriksson, 1994) that 

has not yet been tested using molecular data. Although both 

subfamilies are found in the Andes of Venezuela, only taxa of 

Carludovicoideae are found in Guaramacal. Carludovicoideae 

includes 11 genera and the vast majority of species, whereas 

Cyclanthoideae includes a single, monotypic genus (Cyclanthus

MAP 57. Costus spiralis var. spiralis occurrence in Venezuela.
NUMBER 100 • 5 3

Poit. ex A. Rich.). Characters for distinguishing the two subfamilies are noted in the family description above.

The leaves of Cyclanthaceae could be confused with those 

of Arecaceae, but in Guaramacal the two families are easily 

separated by habit. Our representatives of the former family are 

either hemiepiphytes or epiphytes or ± acaulescent to shortly 

caulescent herbs, whereas our representatives of the latter are 

either shrubby or arborescent and always distinctly caulescent.

KEY TO THE GENERA OF CYCLANTHACEAE

1a. Plants hemiepiphytes, epiphytes or herbaceous lianas; leaves spirally arranged; leaf blades 20–51 cm long . . . . Asplundia

1b. Plants terrestrial; leaves distichous; leaf blades (30–)70–115 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sphaeradenia

Asplundia Harling

Asplundia Harling, Acta Horti Berg. 17: 41. 1954, nom. cons.

Hemiepiphytic, epiphytic or lianescent herbs; acaulescent 

or with short or long stems. Leaves spirally arranged; petioles present, usually equal to or shorter than leaf blades; leaf 

blades bifid (older leaves often secondarily divided in irregular segments), 1- or 3-costate (1-costate in our species), plicate, 

segments oblong, lanceolate or linear-lanceolate, acute or acuminate. Inflorescences axillary, solitary; peduncles short to ~10 

cm long; spathes 3–5(–6), scattered at irregular intervals on peduncles, ovate-lanceolate, subnaviculiform, usually caducous or 

some of the more basal ones persisting; spadix ellipsoid, ovoid 

or cylindrical. Staminate flowers: symmetrical, infundibuliform, 

perianth lobes distributed evenly around the receptacle (our 

species) or asymmetrical with perianth lobes only on one side; 

receptacle flat to deeply concave; perianth lobes glanduliferous; 

stamens few to many. Pistillate flowers: subconnate, separating 

at maturity; tepals distinctly developed; placentae 4, parietal; 

styles 4, ± free or completely concrescent, or lacking; stigmas 

4, variously shaped. Seeds oblong-ovoid, flattened; seed coat 

unsculptured.

A genus of ~100 species found in Mexico, Central America, 

the Lesser Antilles, and South America (Colombia, Venezuela, 

Trinidad and Tobago, the Guianas, Ecuador, Peru, Bolivia, and 

Brazil). Approximately 16 species occur in Venezuela.

KEY TO THE SPECIES OF ASPLUNDIA

1a. Leaf blades 35–51(–90) cm long, usually divided 2⁄3–4

⁄5 their length, segments (3–)4.5–6 cm wide, apices acuminate, 

glabrous above, scattered brown scales sometimes present below . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. moritziana

1b. Leaf blades 20–30 cm long, divided 1⁄2–2⁄3 their length, segments 2.5–4.5 cm wide, apices long-acuminate to caudate, 

glabrous above and below . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. vagans

Asplundia moritziana (Klotzsch) Harling, Acta Horti Berg. 17: 

42. 1954; Harling, in Berry et al., Fl. Venez. Guayana 4: 

474. 1998. Carludovica moritziana Klotzsch, Linnaea 20: 

468. 1847.

Asplundia vagans auct., non (Klotzsch) Harling; Dorr et al., Contr. U.S. Natl. 

Herb. 40: 44. 2000 [2001].

Hemiepiphytes, epiphytes or herbaceous lianas. Petioles (8–)

12–20(–66) cm long; leaf blades 30–51 cm long, bifid (often older 

leaves secondarily cleft in irregular lobes), usually divided 2⁄3–4

⁄5

their length; segments (3–)4.5–6 cm wide, lanceolate, apices acuminate, glabrous above, scattered brown scales sometimes present 

below. Spadix (in fruit) subcylindrical, 4.5–6 × 1–2 cm. Staminate 

flowers: 2 mm long, receptacle distinctly concave; perianth lobes 

obtuse, completely surrounding the receptacle, those on the free 

side of the flower usually larger than the others; stamens 35–50 

or more, almost sessile on “bulbous” bases. Pistillate flowers: 4–6 

mm in diameter, obtusely triangular, equal to or slightly shorter 

than stigmas; stigmas ± laterally compressed, uncinate.

Found in South America (Colombia and Venezuela). In Venezuela, found in the Andes (Lara, Mérida, Portuguesa, Táchira, 

and Trujillo), the Sierra de Perijá (Zulia), the Cordillera de la 

Costa (Aragua, Anzoátegui, Carabobo, Distrito Federal, Falcón, 

Miranda, Monagas, Sucre, and Yaracuy), and the Venezuelan 

MAP 58. Asplundia moritziana occurrence in Venezuela.
5 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Guayana (Amazonas, Bolívar, and Delta Amacuro; Map 58). In 

the park, found in forest understory on both slopes of Guaramacal; 1,500–2,600 m.

Material found in Guaramacal and the Cordillera de 

Mérida appears to be less robust (smaller leaves, narrower leaf 

segments, smaller spadices in fruit, etc.) than collections made in 

the Cordillera de la Costa and in the Venezuelan Guayana, and 

we suspect more than one taxon is subsumed under the name 

Asplundia moritziana. The type of its basionym, Carludovica 

moritziana, is without locality but presumably was gathered in 

Colonia Tovar (Aragua state). If this is the case and there is more 

than one taxon involved, then the name A. moritziana must be 

applied to the material found in the Cordillera de la Costa.

Asplundia vagans Harling, Acta Horti Berg. 18: 170, fig. 45h–

m, t. 16. 1958.

Asplundia sp. A; Dorr et al., Contr. U.S. Natl. Herb. 40: 44. 2000 [2001].

Hemiepiphytic or epiphytic herbs. Petioles 7–14 cm long; 

leaf blades 20–30 cm long, bifid, divided 1⁄2–2⁄3 their length; segments 2.5–4.5 cm wide, linear-lanceolate to lanceolate, apices 

long-acuminate to caudate, glabrous above and below. Spadix (in 

fruit) subcylindrical, 3–4.5 × 1.5 cm. Staminate flowers: not seen. 

Pistillate flowers: 5–6 mm in diameter; tepals triangular, acute to 

acuminate, usually exceeding the stigmas in length, fleshy external 

appendages present; stigmas ± laterally compressed, uncinate.

Found in Central America and South America (Colombia, 

Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, evidently 

restricted to the Andes (Barinas, Portuguesa, and Trujillo; Map 

59). In the park, known only from forest understory on the south 

slope of Guaramacal; (1,200–)1,600 m.

Harling (1958) considered Asplundia vagans to be a widespread but rare species that mainly occurred at lower elevations. 

He noted several subtle differences in the collections he examined. The long-acuminate to caudate apices of the leaf segments, 

however, are distinctive.

Sphaeradenia Harling

Sphaeradenia Harling, Acta Horti Berg. 17: 3. 1954.

Pseudoludovia Harling, Acta Horti Berg. 18: 338. 1958.

Perennial herbs, terrestrial (our species) or epiphytic; stems 

aerial to rhizomatous. Leaves distichous, usually erect; petioles 

distinct, without an adaxial groove or sometimes with a band of 

tissue connecting the margins of the blade with the sheath; leaf 

blades bifid, 1-costate, stiff, thick, coriaceous; leaf segments linear 

to broadly lanceolate or triangular, acute to acuminate, eventually (through environmental factors) divided into several irregular lobes. Peduncles short to long, with a collar-like structure at 

junction with spadix, recurved to straight in fruit. Spathes 2–5, 

ovate to lanceolate, usually on the distal 1⁄2 of a peduncle, usually 

caducous and leaving a discernable scar. Spadix broadly ellipsoid 

to cylindrical, often brightly colored (white, green, yellow or red) 

in fruit. Staminate flowers: asymmetrical, caducous or persisting 

in fruit; receptacles usually flat to slightly concave; perianth lobes 

distributed around the receptacle, abaxial lobes usually larger or 

lobes on abaxial side only, abaxial lobes with tiny glands, adaxial 

lobes usually without tiny glands, lobes obtuse to acute, apices 

straight to curved outward; stamens few to many; anthers with 

or without an apical secretion globule; filaments short to almost 

absent. Pistillate flowers: connate basally; tepals 4, free from each 

other or variously connate, lateral tepals usually broader than 

medial ones; staminodes present, fragrant; ovary embedded in 

the rachis and protruding at maturity; placenta 1, apical; styles 

usually concrescent; stigmas proximally connate to free, laterally 

compressed or not. Seeds terete, broadly ellipsoid to narrowly oblong, straight or curved; testa sculptured.

A genus of ~50 species found in Central America and South 

America (Colombia, Venezuela, Ecuador, Peru, and Brazil). The 

greatest species diversity is found in Colombia. Four species 

occur in Venezuela, of which only one is Andean.

References. Eriksson (1995); Harling (1954).

Sphaeradenia laucheana (Sander ex Mast.) Harling, Acta Hort. 

Berg. 17: 3. 1954; Morillo, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 2: 447–448, fig. 1. 2010. Salmia 

laucheana Sander ex Mast., Gard. Chron., ser. 3, 13: 442. 

[15 Apr] 1893; ibid., 481, fig. 72. [22 Apr] 1893.

The nominate subspecies is the only subspecies found in 

Venezuela.

Sphaeradenia laucheana subsp. laucheana

FIGURE 25B

Plants to 3 m tall; stems 0.2–1.5 m tall, sometimes branched. 

Leaves large, to 2.5 m long; leaf sheaths to 45 cm long; petioles 

10–75 cm long; leaf blades (30–)70–115 cm long, deeply bifid 

(>50% of their length), somewhat coriaceous, segments 4–15 

cm wide, linear to oblong or lanceolate, acuminate (to acute). 

MAP 59. Asplundia vagans occurrence in Venezuela.
NUMBER 100 • 5 5

Peduncles (10–)20–30 cm long; spathes (2–)3–5, 10–15 cm long, 

reddish or white tinged with red, spathe scars visible near middle 

and below on peduncles. Spadix narrowly ellipsoid to cylindrical, green or greenish-white or yellowish-green, 2.5–3 × 1–1.5 

cm (at anthesis), 4–10 × 1.5–4 cm (in fruit). Staminate flowers: 

usually caducous, sometimes persistent, receptacles slightly concave to convex; perianth lobes 9–16, 4–10 lobes on the adaxial 

side, sometimes with tiny glands; stamens numerous (up to 30); 

filaments to 0.3 mm long, densely aggregated. Pistillate flowers: 

tepals ± cuspidate or ± triangular, acute to obtuse or truncate, 

apex incurved and sometimes covering the stigmas; staminodes 

present; ovary scarcely exserted in mature fruit; stigmas connate 

(to free), linear to oblong, lanceolate or narrowly triangular. 

Seeds ellipsoid to oblong, ~4 × 2 mm, curved or unciform.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, known only from the Andes (Barinas, 

Portuguesa, Táchira, and Trujillo; Map 60). In the park, found in 

forest and cloud forest on both slopes of Guaramacal, including 

La Divisoria de la Concepción; 1,600–2,600(–3,000) m.

Eriksson (1995) distinguished two subspecies: the nominate 

one (described above) and Sphaeradenia laucheana subsp. irazuensis (Cufod.) R. Erikss., an epiphytic plant whose staminate 

flowers persist on the mature spadix and whose stigmas are remarkably uncinate in fruit. The latter subspecies is restricted to 

Central America (Costa Rica).

Cyperaceae

M. T. STRONG

(CAREX: M. T. STRONG AND A. A. REZNICEK)

Grass- or rush-like perennial or annual herbs with rhizomes 

or stolons. Culms solitary or caespitose, 3-angled or obtusely so, 

rarely 4- or 5-angled, sometimes terete or flattened, solid or hollow, glabrous or sometimes pubescent. Leaves well developed or 

reduced to bladeless sheaths, borne at base or both basal and cauline; blades flattened, plicate, involute or subcylindric, linear to 

lanceolate, glabrous or sometimes pubescent, usually antrorsely 

scabrous on margins and primary veins or occasionally smooth; 

sheaths closed, often splitting with age, inner band membranous 

with a concave to convex orifice or sometimes entirely herbaceous with the summit frequently prolonged as a triangular or 

tonguelike contraligule (facing away from stem); ligule absent or 

short, sometimes a narrow band of hairs. Inflorescences diverse, 

complex, and difficult to interpret, typically umbelliform, anthelate, corymbose, spicate, paniculate, racemose or capitulate, sessile or branching, bearing 1 to many spikelets or reduced spikes 

from the upper leaflike or sometimes scalelike bracts; spikelets 

1- to many-flowered, often with 1 to several empty (sterile) 

scales at base. Flowers (florets) spirally imbricate or 2-ranked 

(distichous), bisexual or unisexual (the plant then monoecious 

or rarely dioecious), borne singly, each from the axil of a spikelet 

scale; perianth, when present, of 3 to many, smooth or barbed, 

hypogynous bristles or scaly segments, usually persistent at the 

base of the mature achene; stamens 1–3; anthers elliptic to linear, 

apiculate or appendaged apically, basifixed, thecae parallel, longitudinally dehiscent; ovary 2- or 3-carpellate, unilocular, style 

2- or 3-branched. Fruit an achene, trigonous (sometimes dorsally 

compressed), biconvex, globose or cylindric, ellipsoid, oblong or 

obovoid, usually apiculate apically, smooth, reticulate, puncticulate, papillose, verrucose or transversely rugulose, style base 

deciduous or persistent apically.

A cosmopolitan family of ~100 genera and ~5,500 species. 

In Venezuela, 34 genera and ~430 native or naturalized species 

occur in aquatic or terrestrial habitats.

References. Bruhl (1995); Goetghebeur (1998); 

Muasya et al. (1998, 2000, 2009a); Simpson et al. (2007); 

Tucker (1987).

KEY TO THE GENERA OF CYPERACEAE

1a. Inflorescence contracted without evident lateral branches or rays, unispiculate (a solitary spikelet at the summit of the 

culm), capitate cluster of 2 to many spikelets or a linear contiguous spike at the apex of the culm . . . . . . . . . . . . . . . . 2

2a. Inflorescence a linear, simple contiguous spike; fruit an herbaceous sac (perigynium) tightly enclosing a trigonous 

achene and vestigial rachilla bearing a hooked rudimentary spikelet at apex . . . . . . . . . . . . . . . . . . . . . . . Uncinia

2b. Inflorescence unispiculate or a capitate cluster of 2 to many spikelets; fruit a naked achene or if a perigynium, then 

not enclosing a hooked rudimentary spikelet at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

MAP 60. Sphaeradenia laucheana subsp. laucheana occurrence in 

Venezuela.
5 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

 3a. Elongate leaf blades and involucral bracts wanting, summit of the leaf sheaths merely apiculate or entire; inflorescence unispiculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eleocharis

 3b. Elongate leaf blades and/or involucral bracts present; inflorescence multispiculate or unispiculate . . . . . . . . . 4

 4a. Fruit an herbaceous prominently veined sac (perigynium), loosely or tightly enclosing an achene, often 

bidentate apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carex

 4b. Fruit a naked achene, not surrounded by a perigynium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 5a. Achene bearing the persistent expanded base of the style at apex and barbed or plumose bristles at base 

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhynchospora

 5b. Achene merely apiculate or entire at apex, without bristles at base . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 6a. Plants small (<10 cm tall), forming dense cushions or mats in páramo; leaves incurved, stiff, and 

somewhat cartilaginous; flowers with 2 alternating whorls of scalelike tepals at the base that are 

often persistent on rachilla after spikelet scales and achenes have shed . . . . . . . . . . . . . Oreobolus

 6b. Plants medium to large or rarely small (10–200 cm tall), not forming dense cushions or mats and occurring in various habitats; leaves flexuous; flowers lacking scalelike tepals, naked at the base . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperus

1b. Inflorescence open or, if contracted, then with at least some evident lateral branches or rays, terminal or with 2 to several 

lateral partial inflorescences from the upper leaflike bracts, umbelliform, corymbose, paniculate or racemose, 2 to many 

spikelets solitary or in fascicles, digitate or glomerate clusters, or spikes at branch or ray tips . . . . . . . . . . . . . . . . . . . 7

7a. Fruit an herbaceous prominently veined sac (perigynium), loosely or tightly enclosing an achene, often bidentate 

apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carex

7b. Fruit a naked achene, not surrounded by a perigynium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

 8a. Flowers unisexual; achene globose or ovoid, rarely trigonous, with a smooth white, greenish or purple tinged, 

bony or crustaceous pericarp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scleria

 8b. Flowers bisexual; achene trigonous or biconvex with a light brown to brown or blackish puncticulate, cancellate 

or transversely rugulose pericarp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

 9a. Spikelet scales distichous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperus

 9b. Spikelet scales imbricate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

 10a. Style base persistent at achene apex, triangular to lanceolate; style 2-branched or entire; achene biconvex; bristles 1–6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhynchospora

 10b. Style base disarticulating from the achene, achene apex merely apiculate; style 3-branched; achene 

trigonous; bristles absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fimbristylis

Carex L.

Carex L., Sp. Pl. 972. 1753.

Grasslike perennial herbs, tufted or with short or elongate 

creeping rhizomes, sometimes stoloniferous, monoecious, rarely 

dioecious. Culms triquetrous or trigonous, rarely subterete, solid 

or sometimes hollow. Leaves 3-ranked; ligule often present at 

adaxial junction of sheath and blade; blades flattened, V-shaped 

or folded, narrowly linear, rarely lanceolate or elliptic and subpetiolate. Inflorescence a single terminal spike or a series of 1 

to several terminal subcontiguous or remote spikes or panicles 

from the upper leaflike or scalelike bracts; spikes sessile or peduncled with spirally arranged florets (perigynia), unisexual, 

androgynous or gynaecandrous, rarely mixed, inflorescences 

bearing unisexual spikes often with the 1 to several staminate 

spikes terminal and pistillate spikes below. Flowers unisexual; 

hypogynous squamellae or bristles absent; staminate florets with 

3 stamens subtended by a single scale; pistillate florets a single 

ovary contained within a tight or inflated, 2-sided, trigonous or 

rounded, often bottle-shaped perigynium, subtended by a single 

scale; style 2- or 3-branched, continuous with the ovary and persistent or articulated with it and deciduous, straight or flexuous, 

often thickened at base, tips of the stigmas exserted from the 

apex or often bidentate beak of the perigynium; perigynium 

membranous, chartaceous or coriaceous, often ribbed or nerved 

with raised or impressed nerves, or smooth, sometimes winged, 

glabrous, pubescent, hispid, papillose, puncticulate or smooth, 

sometimes spongy at base, apex often beaked, bidentate or bifurcate. Fruit an achene, lenticular, biconvex or trigonous, generally 

green, puncticulate or sometimes smooth.

A cosmopolitan genus of ~2,000 species, widespread in 

temperate zones and montane regions of the tropics; ~30 species 

occur in Venezuela.

References. Gehrke et al. (2010); Hipp et al. 

(2006); Kükenthal (1909); Reznicek (1990); Starr and Ford 

(2009); Starr et al. (2004, 2008); Waterway and Starr (2007); 

Waterway et al. (2009).

Carex is one of the largest genera of vascular plants known, 

and not surprisingly, many taxonomic problems surround its limits. Molecular data (Waterway and Starr, 2007; Starr et al., 2008; 

Starr and Ford, 2009) suggest that Carex as currently construed 

resolves itself into four main clades that are not congruent with 

previous classifications. None of the taxa present in Guaramacal have figured into any of these molecular studies, although 
NUMBER 100 • 5 7

one unresolved problem that affects our flora concerns how one 

deals taxonomically with a number of unispiculate Carex species 

that form a monophyletic clade with Kobresia Willd., Uncinia

Pers. (present in Guaramacal), and Cymophyllus Mack. Until 

this problem is resolved, we have opted to recognize Uncinia as 

a distinct, albeit paraphyletic, genus.

KEY TO THE SPECIES OF CAREX

1a. Style 2-branched; achene biconvex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Spikes 3–4.5 mm wide; perigynia narrowly ovate or lanceolate, rib-margined . . . . . . . . . . . . . . . . . . C. bonplandii

2b. Spikes 4–6 mm wide; perigynia broadly obovate or suborbicular, wing-margined . . . . . . . . . . . . . . . . . . . C. longii

1b. Style 3-branched; achene trigonous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Inflorescence open with nodding spikes; spikes (3–)3.5–8(–10) cm long; achene constricted abaxially . . . C. jamesonii

3b. Inflorescence contracted, with erect to spreading spikes; spikes 0.5–2.5(–3) cm long; achene not constricted . . . . 4

 4a. Terminal spike androgynous; pistillate spikes 2–3 mm wide; perigynia 3.5–4 × 1–1.2 mm . . . . . . . . C. tachirensis

 4b. Terminal spike wholly staminate; pistillate spikes 5–9 mm wide; perigynia 3.7–6 × 1.3–1.7 mm . . . . . C. tamana

Carex bonplandii Kunth, Enum. Pl. 2: 380. 1837; Briceño, in 

Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 456, 

fig. 6. 2010.

Loosely caespitose, (4–)12–60(–80) cm tall; rhizomes short, 

horizontally creeping, culms densely set, arising singly or several together. Culms trigonous, 0.7–1.7 mm wide, smooth 

proximally, antrorsely scabrous on angles distally. Leaves 2–8 

per culm, basal and lower cauline, lowermost often bladeless 

or with short blades, to 30 cm long; sheaths short or elongate, 

light brown to brown, glabrous, inner band membranous, with 

a truncate to slightly concave orifice; ligule a thickened band 

of light brown to brown tissue, acutely oriented; blades linear, 

V-shaped proximally, subflattened-plicate distally, 1–4.5 mm 

wide, attenuate. Inflorescence a small aggregate of 3–13(–16) 

spikes, 1–3.5 × 0.5–1.5(–2) cm; involucral bracts narrowly linear to setaceous, lowermost one 1–4(–8) cm long, often overtopping the inflorescence; spikes gynaecandrous, 6–10 × 3–4.5 mm, 

with 5–30 subappressed to distally spreading perigynia; pistillate 

scales ovate to ovate-lanceolate, acuminate, shallowly cucullate, 

2–3.3 × 1–1.3 mm, slightly shorter than the perigynia but as 

wide, submembranous, pale to dark brown, margins narrowly 

scarious, midcosta wirelike, pale green, shorter than to equaling the slightly emarginate tip. Anthers 1.3–1.7 mm long. Style 

2-branched. Perigynia plano-convex or sometimes concave ventrally, narrowly ovate or lanceolate, 2.8–4 × 1–1.3 mm, thick 

and coriaceous, light green becoming brownish, narrowly wingmargined nearly to the base, faintly or distinctly 7–12-nerved 

dorsally and sometimes ventrally, often nerved ventrally only at 

base, tapering gradually into a shortly bidentate tip, margins antrorsely scabrous above the middle, rib-margined and subinvolute 

proximally. Achenes lenticular, oblong-ovate or ovate-elliptic, 

1.5–2 × 0.7–1 mm, narrowly obtuse apically, subabruptly narrowed to a short-attenuate base, puncticulate, yellowish-brown.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Ecuador, Peru, Bolivia, and Brazil). In 

Venezuela, restricted to the Andes (Mérida, Táchira, and Trujillo; Map 61). In the park, found on both slopes of Guaramacal, 

near the Laguna del Pumar, and in the Páramo de Guaramacal; 

2,000–3,100 m.

Hipp et al. (2006) presented evidence for a possible allopolyploid origin of the Carex bonplandii species complex, which 

consists of C. bonplandii and C. roraimensis Steyerm. The latter 

species is endemic to the Venezuelan Guayana.

Carex jamesonii Boott, Proc. Linn. Soc. Lond. 1: 258. 1845 

[1849]; Reznicek, in Berry et al., Fl. Venez. Guayana 4: 519. 

1998; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 459, fig. 9. 2010.

FIGURES 5, 25C

Coarse, stoloniferous, 55–110(–130) cm tall; rhizomes 

short and knotty, emitting slender, stiff, floccose stolons. Culms 

solitary or several together, triquetrous distally, channeled medially on sides proximally, 1.7–4.8 mm wide, stiff and hardened 

proximally, flexuous distally, smooth on angles proximally, scabrous to smooth on angles distally. Leaves 6–10, primarily basal, 

lowermost often bladeless, 30–120 cm long; sheaths folded and 

MAP 61. Carex bonplandii occurrence in Venezuela.
5 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 5. Cyperaceae, Carex jamesonii Boott. A. Habit. B. Inflorescence. C. Inflorescence spike (detail). D. Spikelet scale. E. Perigynium.

F. Achene dissected from perigynium. (A–F, Stergios et al. 15985.)
NUMBER 100 • 5 9

equitant proximally, brownish-red or red-purple; ligule a broad, 

reddish-brown band of tissue gradually converging from the 

margins to midblade; blades linear, 5–12 mm wide, elongate, 

folded proximally, flattened to subplicate distally, glabrous, 

glaucous, attenuate. Inflorescence a subcontiguous series of 1 

to several nodding spikes in short racemes from the upper leaflike bracts; lowermost 1 or 2 involucral bracts well developed, 

leaflike, uppermost linear to setaceous, their sheaths wanting or 

at most very short; spikes 11–35, androgynous, linear-cylindric, 

(30–)35–80(–100) × 5–7 mm, densely flowered, approximate 

and short peduncled or the lowermost on elongate peduncles 

and often pendulous; staminate portion of spike 10–30 mm long; 

pistillate scales lanceolate, 2.7–4.7 × 1–1.3 mm, nearly equaling 

the perigynia, brown, red-brown or blackish, narrowly scarious, 

midcosta 3-nerved, green or pale brown, prolonged beyond the 

acute to acuminate apex as a short setose-scabrous awn. Anthers 

1–2 mm long. Style 3-branched. Perigynia ellipsoid-lanceoloid, 

compressed trigonous, often falcate when mature, (2.7–)3–4.3 × 

1–1.5 mm, gradually or abruptly attenuate to a shallowly bidentate apex, cuneate to short-stipitate base, conspicuously nerved 

with 2 coarse marginal ribs, pale to red-brown or blackish, glabrous or scabrous distally, lustrous. Achenes trigonous, oblongellipsoid or oblong-obovoid, 1.5–2.5 × 0.7–1 mm, obtuse to 

acute at both ends, abaxial face strongly constricted or wrinkled 

medially, puncticulate, pale brown.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, found in the Andes (Apure, Lara, Mérida, Táchira, and 

Trujillo), the Cordillera de la Costa (Anzoátegui, Aragua, Distrito Federal, Miranda, Monagas, Sucre, and Yaracuy), and the 

Venezuelan Guayana (Bolívar; Map 62). Found in cloud forest, 

subpáramo, and páramo in a number of localities throughout the 

park; (1,500–)2,000–3,100 m.

The inflorescences of several collections made in the park 

(e.g., Cuello et al. 1072, 1359; Niño et al. 1359) are infected 

with a smut.

• Carex longii Mack., Bull. Torrey Bot. Club 49: 373. 1922 

[1923]; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 461–462. 2010.

Caespitose, forming small to medium-sized clumps, (15–)

30–80(–120) cm tall; rhizomes short. Culms erect, slender, 1.3–3 

mm wide, trigonous to sharply so, stiff and hardened, smooth 

proximally, scabrous on angles at apex. Leaves 4–6, lower cauline 6–40 cm long, basal ones often bladeless; sheaths elongate, 

tight around culm, proximal ones pale brown, cauline ones 

green, inner band herbaceous, prominently veined, with concave 

orifice; ligule rounded, 2–6 mm long; blades linear, 1.5–4(–4.5) 

mm wide, plicate, glabrous, long-acuminate. Inflorescence a 

series of (2–)3–10(–11), subcontiguous, sessile, ellipsoid to obovoid spikes at the summit of the culm, (1.4–)2.2–6 cm long; 

inflorescence bracts setaceous, reduced distally; spikes gynaecandrous, 6–17 × 4–6 mm, obtuse, scales appressed-ascending or 

slightly spreading; pistillate scales ovate, 2.2–3.7 × 1.1–1.8 mm, 

obtuse to acute, lustrous, silvery white to pale brown, midcosta 

3-nerved, green, midnerve prolonged as a short awn or mucro 

on proximal scales, ending short of the tip on distal scales. Anthers 1–2.3 mm long. Style 2-branched. Perigynia plano-convex, 

broadly obovate or suborbicular, broadest near the middle, 

3–4.5 × 1.6–2.8 mm, light silvery-green or silvery-brown, 

MAP 62. Carex jamesonii occurrence in Venezuela. MAP 63. Carex longii occurrence in Venezuela.
6 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

3–9-nerved adaxially, 5–13-nerved abaxially, margins thinly 

winged, abruptly contracted into a scabrous-margined, bidentate 

beak, 0.7–1.3 mm long, with teeth ~0.2–0.4 mm long. Achenes 

biconvex, ovate to oblong-ovate, 1.4–1.7 × 0.9–1.1 mm, shortstipitate, pale brown.

Native to North America; adventive in Mexico, Central 

America, the West Indies, South America (Colombia, Venezuela, Ecuador, Brazil, Argentina, and Uruguay), the Pacific Islands 

(Hawaii), and New Zealand. All Venezuelan records are from 

the Andes (Lara and Trujillo; Map 63). In the park, found on 

both slopes of Guaramacal; 1,900–2,800 m.

Carex tachirensis Steyerm., Fieldiana, Bot. 28: 68, fig. 8I–M. 

1951.

Rhizomatous, 3–34 cm tall; rhizomes ascending to horizontal, slender or knotty, clothed with fibrous sheaths. Culms sharply 

ascending, trigonous, often channeled along 1 margin, antrorsely 

spinulose-scabrous on angles and often on sides (at least distally). Leaves primarily basal, 2–11 cm long; sheaths short; ligule 

a narrow band of tissue at adaxial junction of sheath and blade; 

blades linear-lanceolate, 1.3–3.5 mm wide, stiffly erect, rigid, plicate, attenuate. Inflorescence racemose, of a terminal and 1 or 2 

lateral long-pedunculate spikes; involucral bracts 1(2) per spike, 

leaflike, sheathing; spikes narrowly linear, terminal one androgynous, lateral wholly pistillate, 10–15 × 2–3 mm; pistillate scales 

widely ovate or lanceolate, 3.5–6 × 1.5–2 mm, castaneous or 

dark reddish-brown on sides, margins entire, midcosta broad, 

pale green, 3-nerved with spinulose-scabrous to smooth nerves, 

prolonged beyond the short-acuminate apex as an elongated setulose awn on lower scales, upper scales short-awned; staminate 

scales ovate, acuminate, ~3 mm long, like the pistillate scales. 

Anthers ~1 mm long; style 3-branched. Perigynia obscurely trigonous, elliptic-lanceolate, 3.5–4 × 1–1.2 mm, 3-angled, thinly 

herbaceous, 3–5-nerved on each side, short-stipitate at base, narrowing distally to a slender, setulose, bidentate beak. Achenes 

trigonous with the face against the rachilla concave, elliptic to 

oblong-elliptic or elliptic-obovate, ~2 × 1 mm, obtuse apically, 

cuneate at base, puncticulate, brown.

Endemic to the Andes of Venezuela (Táchira and Trujillo; 

Map 64). Páramo near the Laguna del Pumar; 3,000 m.

We have a single collection (Niño et al. 1515, PORT) of this 

species from the park but suspect that Carex tachirensis is more 

common and overlooked by collectors.

Carex tamana Steyerm., Fieldiana, Bot. 28: 70. 1951; Reznicek, 

in Berry et al., Fl. Venez. Guayana 4: 519. 1998.

Carex acutata auct., non Boott; Ortega et al., BioLlania 5: 40. 1987.

Loosely caespitose to mat-forming, 8–60(–80) cm tall; rhizomes often emitting scaly, cord-like stolons. Culms ascending to 

erect, trigonous to obtusely so proximally, 1–2.2 mm wide, hardened but subflexuous, smooth, angles bluntly and closely antrorsely scabrous. Leaves 7–12, basal and lower cauline, sharply 

ascending-spreading, 2–35 cm long; sheaths short, herbaceous, 

scabridulous, pale brown to brown proximally, inner band membranous, scabridulous, light reddish-brown, orifice concave on 

proximal sheaths, truncate to convex on distal ones; ligule a 

submembranous band of tissue acutely oriented; blades thickly 

herbaceous, subrigid, flattened-plicate, folded to V-shaped proximally, 2.2–4.4 mm wide, scabrous adaxially, smooth abaxially, 

long-acuminate. Inflorescence a terminal, compact, spicate cluster 

MAP 64. Carex tachirensis occurrence in Venezuela. MAP 65. Carex tamana occurrence in Venezuela.
NUMBER 100 • 6 1

of 2–4 pistillate spikes and a single terminal staminate spike, 

often bearing below a remote lateral pistillate spike on a short 

peduncle; involucral bracts leaflike, strongly ascending, essentially sheathless, each subtending the base of a spike, overtopping 

the spikes but becoming successively shorter distally; pistillate 

spikes erect to spreading, oblong-cylindric or oblong-obovoid, 

5–25(–30) × 5–9 mm, perigynia often spreading; staminate 

spike linear, 1.6–2.7 cm long; pistillate scales ovate-lanceolate to 

elliptic-lanceolate or lanceolate, glabrous, light brown, margins 

narrowly scarious medially, midcosta green, slender and finely 

3-nerved, lateral nerves indistinct, medial nerve distinct, ending short of or shortly prolonged beyond the narrowly acute to 

acuminate apex; staminate scales linear, ~8 mm long. Anthers 

1.5–2 mm long. Style 3-branched. Perigynia trigonous, lanceoloid, 3.7–6 × 1.3–1.7 mm, adaxial side plane, abaxial sides plane 

or somewhat swollen, slightly contracted above the middle to a 

slender, acuminate to long-acuminate beak that is bidentate apically, obtuse to subrounded at base, short-stipitate, distinctly and 

finely nerved with several coarse nerves, glossy, green to brownish. Achenes trigonous, ellipsoid, with plane or slightly concave 

sides, ~2 mm long, puncticulate, light brown.

Found in Central America (Costa Rica) and South America 

(Colombia, Venezuela, and Ecuador). In Venezuela, this species 

has a disjunct distribution occurring in the Andes (Táchira and 

Trujillo) and in the Venezuelan Guayana (Bolívar; Map 65). In 

the park, found near the Laguna del Pumar and in the Páramo de 

Guaramacal; (2,500–)3,000 m.

Cyperus L.

Cyperus L., Sp. Pl. 44. 1753.

Kyllinga Rottb., Descr. Icon. Rar. Pl. 12. 1773, nom. cons.

Pycreus P. Beauv., Fl. Oware 2: 48. 1816.

Lipocarpha R. Br., in Tuckey, Narr. Exped. Zaire 459. 1818, nom. cons.

Ascolepis Nees, in Steudel, Syn. Pl. Glumac. 2: 105. 1855, nom. cons.

Grasslike perennial or annual herbs, glabrous; rhizomes, 

when present, short or stoloniferous, horizontally creeping, 

elongate or short knotty, or tufted. Culms trigonous or sometimes terete, smooth or scabrous, green. Leaves primarily basal 

with several often lower cauline; sheaths finely veined, sometimes cross-veined, usually glabrous; ligule absent or short; 

blades herbaceous or stiffened, flattened, V-shaped, plicate, 

inrolled, terete or crescentform, finely veined abaxially, finely 

cellular-reticulate and semiglossy adaxially, sometimes septate 

or cross-veined, margins, abaxial midvein, and adaxial lateral 

veins, when present, usually antrorsely scabrous. Inflorescence a 

simple or compound, terminal, umbel-like corymb with spreading rays, rarely congested or pseudolateral and head-like, or 

subglobose head of 1–3(–4) crowded, sessile spikes; spikes cylindric, ovoid or globose, densely flowered with slender rachis; 

involucral bracts generally leaflike, approximate, spreading in a 

radius; spikelets ovate, ovate-lanceolate or linear in shape, flattened, compressed or subcompressed, subcylindric or 4-angled 

in cross section, palmately, pinnately or imbricately arranged 

on the rachis, many-flowered, lowermost scale and subtending 

bract empty; scales 2-ranked, ovate, oblong-ovate or elliptic, 

acutely keeled, naviculiform or subrounded in cross section, 

sometimes 2-keeled, often longitudinally veined on sides, with a 

1–5-nerved often greenish midcosta, apex acute, obtuse or cuspidate, often mucronulate; rachilla winged from the persistent, 

decurrent base of the scale or wingless, wings hyaline or sometimes corky or thickened, disarticulating at base from the rachis, 

whole spikelet falling entire or disarticulating at the nodes or 

base, internodes, scales, and achenes falling as 1-fruited segments or rachilla persistent on the rachis, scales and achenes 

deciduous. Flowers bisexual or staminate; hypogynous squamellae or bristles absent; stamens 1–3; anthers oblong-elliptic, 

narrowly elliptic or linear, apiculate or sometimes minutely appendaged apically. Styles 2- or 3-branched. Achenes trigonous 

or lenticular, ovoid, obovoid, oblong-obovoid, ellipsoid or narrowly ellipsoid, or usually short-apiculate apically, sometimes 

shortly stipitate at base, surface puncticulate, reticulate or 

smooth, rarely transversely wrinkled, or achenes laterally flattened with 1 angle facing the rachilla, narrowly ovate to oblong or elliptic, obtuse to subrounded or subtruncate apically, 

apiculate, cuneate to rounded at base, substipitate to distinctly 

stipitate, surface puncticulate.

A cosmopolitan genus of 600–700 species found in temperate, subtropical, and tropical regions. In the Americas, Cyperus

is found in North America (Canada and USA), Mexico, Central 

America, the West Indies, and South America (all country-level 

political units). The genus evidently is adventive in Canada. Approximately 77 species occur in Venezuela.

References. Bauters et al. (2014); Corcoran (1941); 

Kükenthal (1935–1936); Larridon et al. (2011, 2013, 2014); 

Muasya et al. (1998, 2001, 2002, 2009b); Tucker (1984, 1994).

Cyperus L. is the second largest genus in the Cyperaceae, 

principally tropical, and widely distributed. However, molecular data suggest that Cyperus as presently construed is 

not monophyletic as many other smaller cyperoid genera that 

are currently recognized are embedded within it. Cyperus and 

nine smaller genera form a strongly supported clade with two 

well-supported subclades: a eucyperoid clade associated with 

C3

 photosynthesis and a chlorocyperoid clade associated with 

C4

 photosynthesis. Pycreus P. Beauv. and Kyllinga Rottb. are 

deeply embedded in the latter subclade (Muasya et al., 1998, 

2001, 2002, 2009b; Larridon et al., 2011, 2013, 2014), which 

also contains the nomenclatural type of the genus Cyperus. Despite strong evidence for these relationships, there has been reluctance to place Pycreus and especially Kyllinga in synonymy. 

We have chosen to fold these smaller genera into Cyperus and 

accept a large genus with two infrageneric groups based on C3

and C4

 anatomy.

Although three of the generic synonyms of Cyperus listed 

above are conserved names, none of these names is conserved 

against Cyperus.
6 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

KEY TO THE SPECIES OF CYPERUS

1a. Spikelets 1(2)-flowered; rachilla short; scales conduplicate, conspicuously keeled . . . . . . . . . . . . . . . . . . . . . C. hortensis

1b. Spikelets 2- to many-flowered; rachilla elongate; scales navicular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Style 3-branched; achenes trigonous with a face against the rachilla; spikelet scales yellow-brown; rachilla narrowly 

winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. hermaphroditus

2b. Style 2-branched; achenes biconvex with an edge against the rachilla; spikelet scales dark red or dark reddish-brown; 

rachilla not winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. niger

Cyperus hermaphroditus (Jacq.) Standl., Contr. U.S. Natl. Herb. 

18(3): 88. 1916; Tucker, in Berry et al., Fl. Venez. Guayana 

4: 530. 1998. Carex hermaphrodita Jacq., Collectanea 4: 

174. 1790 [1791]. Mariscus hermaphroditus (Jacq.) Urb., 

Symb. Antill. 2(2): 165. 1900.

Rhizomatous, glabrous perennial, 15–90 cm tall; rhizomes 

short, horizontal, bearing culms singly or sometimes 2 or 3 together. Culms erect, often tuberous-thickened at base, triquetrous, stiff and hardened, 1–3 mm wide, smooth, green. Leaves 

3–10, primarily basal, 15–80 cm long; sheaths eligulate, light 

reddish-brown or purple tinged, inner band membranous, concave at orifice; blades linear, 2–11 mm wide, V-shaped to subflattened, attenuate apically, generally shorter than the culm. 

Inflorescence a simple or sometimes partially compound umbellike corymb with ascending rays, 3–14 × 5–10 cm; involucral 

bracts leaflike, 3–8, lowermost 2–4 elongate, spreading, overtopping the inflorescence; rays 3–12, unequal, to 12 cm long, each 

bearing 1(–3) spikes at tips; spikes loosely to subdensely oblongcylindric, 1–4.5 × 0.7–2.2 cm, obtuse apically; spikelets linear to 

narrowly ellipsoid, 4–9 × 1–1.2 mm, quadrate or subterete, divaricately spreading, lowermost spikelets on spike often reflexed, 

narrowly acute apically, cuneate at base, with 3–8 scales; rachilla 

deciduous, narrowly winged; scales only slightly overlapping the 

base of the ones above, strongly ascending to appressed, ovateelliptic, 2.5–3.3 × 1.2–2 mm, submembranous, pale greenish to 

golden- or reddish-brown, slenderly 3–4-nerved on sides, with 

narrow scarious margins, midcosta green, 3-nerved, ending just 

below the obtuse apex or prolonged as a short mucro. Stamens 

3; anthers 0.5–1 mm long. Style 3-branched. Achenes trigonous 

with flat or slightly concave faces, oblong-ellipsoid to oblongobovoid, 1.2–1.9 × 0.6–0.9 mm, slightly apiculate, estipitate, 

puncticulate, light brown or reddish-brown at maturity.

Found in North America (USA), Mexico, Central America, 

the West Indies, and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Paraguay, and Argentina). Widely 

distributed in Venezuela (Anzoátegui, Apure, Aragua, Bolívar, 

Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, 

Monagas, Táchira, Trujillo, and Zulia; Map 66). Evidently recently adventive in the park, where it has been collected in wet, 

disturbed areas on the north slope along the Boconó–Guaramacal road and on the south slope near El Campamento below 

Cerro El Diablo; ~1,800–2,000 m.

Cyperus hortensis (Salzm. ex Steud.) Dorr, comb. nov. Kyllinga 

hortensis Salzm. ex Steud., Syn. Pl. Glumac. 2: 68. 1854.

Kyllinga pumila Michx., Fl. Bor.-Amer. 1: 28. 1803; Tucker, in Berry et al., 

Fl. Venez. Guayana 4: 590. 1998; Dorr et al., Contr. U.S. Natl. Herb. 

40: 45. 2000 [2001]; non Cyperus pumilus L., Cent. Pl. II. 6. 1756. 

Cyperus densicaespitosus Mattf. & Kük., in Engler, Pflanzenr. IV, 20 

(Heft 101): 597. 1936, nom. nov.

Kyllinga viridiflora Steud., Nomencl., 2nd ed., 1: 852. 1840, nom. nud.

Densely tufted annual or short-lived perennial, 7–40(–65) 

cm tall. Culms firm but flexuous, triquetrous distally, often trigonous near base, 0.7–1.3 mm wide. Leaves 3–5, basal ones often 

bladeless; sheaths elongate, basal ones pink tinged or reddish 

with dark red veins; blades 4–20(–30) cm × 1.5–3(–3.8) mm, 

flattened to V-shaped. Involucral bracts 3–5, leaflike, slightly ascending to horizontal, rarely the lowermost erect; spikes 1–3, 

globose-ovoid to cylindric, terminal one 5–8(–10) × 4–8 mm, 

lateral secondary spikes smaller; spikelets 50–150, lanceolate to 

oblong-lanceolate, (1.9–)2.3–3(–3.8) × 0.7–1.1 mm; fertile scales 

ovate, 1.8–3.1(–3.4) × 1–1.7 mm, pale brownish to hyaline, laterally 2–4-nerved, keel green, unwinged, with 3–10 antrorsely 

spinulose-scabrous barbs, up to 0.2 mm long, rarely smooth, 

prolonged apically as a short mucro. Stamens 2; anthers 0.2–0.4 

mm long. Achenes ellipsoid-obovoid to oblong-obovoid, 1–1.2

(–1.4) × 0.5–0.7 mm, subtruncate apically, cuneate to subrounded at base, light brown to yellow-brown.

MAP 66. Cyperus hermaphroditus occurrence in Venezuela.
NUMBER 100 • 6 3

A widespread species found in North America (USA), 

Mexico, Central America, the West Indies, South America (all 

country-level political units except Chile), and tropical Africa. It 

occurs throughout Venezuela (Amazonas, Apure, Aragua, Barinas, Bolívar, Cojedes, Delta Amacuro, Distrito Federal, Falcón, 

Guárico, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, 

Táchira, Trujillo, Yaracuy, and Zulia; Map 67). Disturbed wet 

areas of montane forest on both slopes of Guaramacal; 1,900–

2,100(–2,600) m.

This species has long been included in Kyllinga, which was 

traditionally circumscribed as including species with head-like 

inflorescences, deciduous spikelets, and laterally flattened gynoecia. Although the species of Kyllinga form a strongly supported 

monophyletic clade (Larridon et al., 2013, 2014), the clade is 

nested within a larger Cyperus clade composed of species with 

a C4

 photosynthetic pathway. Furthermore, the morphological 

characters that were thought to define Kyllinga are not unique; 

capitate inflorescences with reduced, deciduous spikelets are 

found in various lineages in the larger C4 Cyperus clade, and 

laterally flattened gynoecia occur in both Pycreus and the monotypic Queenslandiella Domin. Therefore, there is little reason to 

support Kyllinga.

Cyperus pumilus L. prevents a combination in Cyperus

based on Kyllinga pumila, and the earliest available epithet 

appears to be the one chosen here. Kyllinga viridiflora Roxb. 

ex Spreng. generally is considered to be a synonym, but references to its publication in a seed list are indirect and imprecise 

bibliographically, and the indirect references lack validating descriptions. If it can be established that K. viridiflora was validly 

published in a seed list, a combination based on its epithet would 

be the correct name for K. pumila in Cyperus.

Cyperus niger Ruiz & Pav., Fl. Peruv. 1: 47. 1798; Briceño, in 

Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 464, 

fig. 14. 2010. Pycreus niger (Ruiz & Pav.) Cufod., Bull. 

Jard. Bot. Belg. 40: 1443. 1970.

FIGURE 6

Rhizomatous perennial, 7–30 cm tall; rhizomes short, slender, ascending. Culms solitary or in small clumps of 2 or 3, 

sometimes decumbent and rooting at lower nodes, triquetrous, 

smooth, easily compressed. Leaves 3–6, primarily basal, 1–3 

cauline, 3–12 cm long; sheaths eligulate, cauline elongate, green 

abaxially, inner band brown or reddish-brown; blades linear, 

1.2–2.5 mm wide, flattened, attenuate apically, generally shorter 

than the culm. Inflorescence contracted and lobate-capitate or 

a small umbel-like corymb with ascending rays, 0.6–3.5 cm in 

diameter, with (1–)3–20 spikelets; involucral bracts leaflike, generally 3, lowermost elongate, ascending to divergent, overtopping the inflorescence; rays 1–4 or absent, when present, short, 

bearing small clusters of 1–3 spikelets at tips; spikelets ovatelanceolate to oblong-lanceolate, 3.5–8.5 × 2–2.3 mm, laterally 

compressed, bluntly acute, 7–18-flowered; rachilla not winged; 

scales densely distichous, ovate to broadly so, 1.4–2 × 1.3–1.8 

mm, dark red or dark reddish-brown, lustrous, smooth on sides, 

midcosta green, 3-nerved, not prolonged beyond the obtuse 

apex. Stamens 2; anthers 0.4–0.5 mm long. Style 2-branched. 

Achenes biconvex, elliptic to broadly elliptic-obovate, 1–1.2 × 

0.7–0.8 mm, very finely and indistinctly transversely wrinkled, 

densely puncticulate, obtuse or rounded to broadly so apically, 

apiculate, short-cuneate at base, sublustrous, brown or blackish.

Found in North America (USA), Mexico, Central America, 

South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, 

Brazil, Chile, and Argentina), and tropical Africa. In Venezuela, 

MAP 67. Cyperus hortensis occurrence in Venezuela. MAP 68. Cyperus niger occurrence in Venezuela.
6 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

known only from the Andes (Barinas, Mérida, Táchira, and Trujillo; Map 68). In the park, found in disturbed areas near montane forest on both slopes of Guaramacal; 1,800–2,000 m.

This species is sometimes assigned to the segregate genus 

Pycreus, but as discussed by Larridon et al. (2013, 2014), Pycreus is not monophyletic, and the presence of laterally flattened 

gynoecia is not sufficient to maintain Pycreus as a genus distinct 

from Cyperus s.l.

Eleocharis R. Br.

Eleocharis R. Br., Prodr. 224. 1810.

Websteria S. H. Wright, Bull. Torrey Bot. Club 14: 135. 1887.

Chillania Roiv., Ann. Bot. Soc. Zool.-Bot. Fenn. “Vanamo” 4(7): 2. 1933.

Egleria L. T. Eiten, Phytologia 9: 481. 1964.

Perennial or occasionally annual herbs, rhizomatous, stoloniferous or tufted; culms usually cylindric, sometimes trigonous, 

FIGURE 6. Cyperaceae, Cyperus niger Ruiz & Pav. A. Habit. B. Inflorescence. C. Spikelet (detail). D. Spikelet scale. E. Flower. F. Achene. (A–C, 

E, Stergios et al. 20718; D, F, Stergios et al. 19321.)
NUMBER 100 • 6 5

rarely quadrangular, hollow or solid, sometimes transversely 

septate, smooth, green, glabrous. Leaves reduced to bladeless 

sheaths; sheaths tubular, orifice truncate to obliquely truncate, 

dorsally carinate, carina often prolonged as a short apiculus, 

sometimes with a puckered, scarious appendage above the apex, 

glabrous. Inflorescence a single, terminal spikelet at the summit of the culm; spikelets ovoid to lanceoloid, rarely obovoid, 

obtuse to acute, several- to many-flowered, with spirally imbricate or sometimes subdistichous scales, lowermost 1 or 2 sterile; 

scales oblong, oblong-ovate to lanceolate or obovate, sometimes 

subrounded, firm or membranous, usually with hyaline margins, carinate or sometimes indistinctly so, nerved or nerveless 

on sides, glabrous. Flowers bisexual; perianth bristles present 

or rarely absent, (3–)6(–12), retrorsely barbed, rarely smooth; 

stamens (2–)3; anthers elliptic to linear. Styles capillary, 2- or 

3-branched, with expanded base. Achenes trigonous or biconvex, obovate or elliptic, smooth, cancellate or cellular-reticulate; 

style base expanded, triangular or conical, sometimes spongythickened, persistent at the apex of the achene.

A cosmopolitan genus of ~250 species with one of its centers of diversity in tropical America. The genus is found in all 

country-level political units in the Americas except Greenland; 

~40 species occur in Venezuela.

References. Hinchliff et al. (2010); Roalson and 

Friar (2000); Roalson et al. (2010); Svenson (1929, 1939).

KEY TO THE SPECIES OF ELEOCHARIS

1a. Achenes terete or essentially so, surface with prominent longitudinal ridges separated by numerous slender trabeculae 

(cross ridges); style base linear-lanceolate, often falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. stenocarpa

1b. Achenes biconvex (2-sided), surface without prominent longitudinal ridges, smooth to finely cancellate or pittedreticulate; style base conic or deltate to depressed-deltate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Culms lacking septate partitions; orifice of the leaf sheath with a conspicuous, often wrinkled or puckered, scarious 

appendage; style base conic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. flavescens

2b. Culms with septate partitions (sometimes indistinct); orifice of the leaf sheath entire, lacking a scarious appendage, 

oblique or truncate, with a dark brown rim; style base deltate to depressed-deltate . . . . . . . . . . . . . . . E. montana

Eleocharis flavescens (Poir.) Urb., Symb. Antill. 4(1): 116. 1903 

(“Heleocharis”); González-Elizondo and Reznicek, in Berry 

et al., Fl. Venez. Guayana 4: 554. 1998. Scirpus flavescens

Poir., in Lamarck, Encycl. 6(2): 756. 1805.

Stoloniferous perennial, (1–)4–40(–50) cm tall; rhizomes 

horizontal to ascending, 0.6–1.2 mm thick. Culms caespitose or spaced singly along the rhizome, erect to ascending, 

subcompressed, 0.4–1 mm wide, prominently ribbed. Sheaths 

subloose, coarsely veined, membranous, green, red tinged near 

base, with a prolonged, inflated, puckered, scarious appendage above the apex, apex short-mucronate dorsally. Spikelets 

ovoid to ovoid-ellipsoid, 3–7 × 1.5–3 mm, subacute, with many 

spirally imbricate florets; fertile scales naviculiform, elliptic to 

narrowly elliptic, 1.2–2.1 × 0.8–1.1 mm, broadly obtuse dorsally, pale green to stramineous, midcostal region green, narrow, 3–5-nerved, subthickened at tip, ending short of the obtuse 

to subacute apex. Stamens 3; anthers 0.7–1 mm long. Styles 

2-branched. Achenes biconvex, obovate, 0.8–1 mm long (including style base), 0.6–0.7 mm wide, smooth to finely cancellate, 

lustrous, olive-brown to dark reddish-brown; style base conic, 

0.2–0.3 × 0.2 mm at base, light brown; bristles 6–8, rarely absent, whitish to tawny, retrorsely barbed, equaling or exceeding 

the style base.

Found in North America (USA), Mexico, Central America, 

the West Indies, and South America (Colombia, Venezuela, 

Trinidad and Tobago, the Guianas, Bolivia, Brazil, Chile, Paraguay, Argentina, and Uruguay). In Venezuela, found in the Andes 

(Lara, Mérida, Portuguesa, Táchira, and Trujillo), the Cordillera de la Costa (Anzoátegui, Aragua, Distrito Federal, Falcón, 

Guárico, Miranda, and Sucre), and the Venezuelan Guayana 

(Amazonas, Bolívar, and Delta Amacuro; Map 69). In the park, 

found in wet areas near montane forest on the south slope of 

Guaramacal; (1,300–)1,800–2,000 m.

Eleocharis montana (Kunth) Roem. & Schult., Syst. Veg. 2: 153. 

1817; González-Elizondo and Reznicek, in Berry et al., Fl. 

MAP 69. Eleocharis flavescens occurrence in Venezuela.
6 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY
NUMBER 100 • 6 7

Venez. Guayana 4: 557. 1998. Scirpus montanus Kunth, in 

H. B. K., Nov. Gen. Sp. [fol.] 1: 182. 1815 [1816]; ibid. 

[qu.] 1: 226. 1815 [1816].

FIGURE 7

Rhizomatous perennial, (20–)30–80(–120) cm tall; rhizomes decumbent, stout, 4–8 mm thick. Culms erect, cylindric, 

striate, conspicuously transversely septate or sometimes inconspicuously so, 1–2.5 mm wide. Sheaths elongate, finely striate, 

purplish-red proximally, stramineous or purple tinged distally, 

red-dotted, orifice truncate with blackish maculate horizontal 

band and a toothlike mucro dorsally. Spikelets ovoid-lanceoloid 

to oblong-lanceoloid, 7–15(–24) × 2–4 mm, acuminate or rarely 

obtuse, many-flowered, florets spirally imbricate; fertile scales 

ovate-lanceolate, (1.5–)2–2.5 × 0.8–1.7 mm, light brown, often 

dark brown medially, reddish-lineolate, margins broadly scarious, midcosta slender, 1-nerved, indistinct, ending short of the 

obtuse or rounded, slightly dilated apex; sterile basal scale 1, 

subrounded, herbaceous, margins broadly scarious. Stamens 3; 

anthers 0.8–1 mm long. Styles 2- or 3-branched. Achenes biconvex, obovate to broadly obovate, 0.9–1.1 × 0.6–1 mm, distinctly 

and finely pitted-reticulate, yellow to brown or olivaceous; style 

base flattened, deltate to depressed-deltate, 0.3–0.4 × 0.4–0.7 

mm at base, shallowly articulated with apex of achene body; 

bristles 4–6, unequal, minutely retrorsely spinulose or somewhat 

smooth, equaling or slightly longer than achene body, ferrugineous or brownish.

Found in North America (USA), Mexico, Central America, 

the West Indies, and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Chile, Paraguay, Argentina, and 

Uruguay). Venezuelan collections are from the Andes (Barinas, 

Lara, Mérida, Portuguesa, Táchira, and Trujillo), the Cordillera 

de la Costa (Anzoátegui, Aragua, Carabobo, Distrito Federal, 

Falcón, Miranda, and Yaracuy), and the Venezuelan Guayana 

(Bolívar; Map 70). In the park, found in wet areas on both slopes 

of Guaramacal; 1,700–2,800 m.

Eleocharis stenocarpa Svenson, Rhodora 31: 205, fig. 35. 1929; 

Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 2: 467. 2010.

Rhizomatous perennial, 6–50 cm tall; rhizomes horizontally 

creeping, scaly, 1–2 mm thick. Culms borne in small clumps along 

rhizome, erect to ascending, subrigid, finely ribbed and channeled, 

green; sheaths elongate, green distally, dark brown or purple-brown 

basally, remotely transversely septate, membranous distally, oblique 

at orifice, dorsally oblong-acuminate with a slender green midcosta 

ending short of the narrowly obtuse to acute apex. Spikelets narrowly ovoid-ellipsoid, 5–9 × 1.4–2.3 mm, acute, 4–16-flowered, 

florets spirally imbricate, scales spreading to divergent with developing achenes; scales narrowly ovate-lanceolate to lanceolate or 

oblong-lanceolate, acuminate, 3–4.5 × 0.9–1.5 mm, subscarious 

and translucent on sides, midcostal region green, 3–5-nerved, ending short of the blunt apex. Stamens 3; anthers 0.8–1.2 mm long. 

FIGURE 7. (Facing page) Cyperaceae, Eleocharis montana (Kunth) Roem. & Schult. A. Habit. B. Inflorescence (detail). C. Spikelet scale.

D. Flower. E. Achene. (A, E, Stergios & Caracas 19308; B–D, Stergios & Caracas 18989.)

MAP 70. Eleocharis montana occurrence in Venezuela. MAP 71. Eleocharis stenocarpa occurrence in Venezuela.
6 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Styles 3-branched. Achenes narrowly obovoid, 1.4–1.8 × 0.5–0.7 

mm, longitudinally costate, very finely and closely trabeculate; style 

base linear-lanceolate, often falcate, often subabruptly expanded at 

base, 0.7–1.2 × 0.2–0.3 mm at base; bristles 2–4, coarse, whitish, 

retrorsely barbed, overtopping the achene.

Found in South America (Colombia, Venezuela, Ecuador, 

Bolivia, and Brazil). Venezuelan material is from the Andes 

(Mérida, Táchira, and Trujillo; Map 71). In the park, found in 

wet open areas in montane forest and páramo on both slopes of 

Guaramacal and in the Páramo de Guaramacal; 1,600–3,100 m.

Common Name. Junco.

Use. This species is used to treat kidney and liver ailments (R. Caracas, pers. comm.).

Fimbristylis Vahl

Fimbristylis Vahl, Enum. Pl. 2: 285. 1806, nom. cons.

Perennial or annual herbs, rhizomatous or tufted with fibrous 

roots. Culms cylindric or subcompressed to flattened, finely to 

coarsely ribbed, smooth, glabrous. Leaves basal; sheaths glabrous 

or pubescent, ventrally splitting with age, apex entire; ligule absent 

or present; blades flattened or inrolled, sometimes folded, narrowly linear to filiform, glabrous or pubescent, margins smooth 

or scabrous. Inflorescence an open, simple or compound anthela 

with ascending rays or sometimes a head-like cluster with congested spikelets; involucral bracts leaflike, usually smaller than the 

leaves, rarely overtopping the inflorescence; primary and secondary rays (when present) unequal, cylindric to subflattened, smooth 

or scabrous; spikelets many-flowered, ovoid, oblong-ovoid, subglobose or lanceoloid, borne singly at ray tips or sometimes clustered; scales spirally imbricate, occasionally somewhat distichous, 

all fertile except for the lowermost one, ovate to oblong-ovate, obtuse to acute, glabrous or pubescent, 1–5-nerved on sides, dorsally 

with an often 3-nerved midcosta, nerves converging apically and 

forming a mucro. Flowers bisexual; hypogynous squamellae or 

bristles absent. Stamens 1–3; anthers oblong or linear, sometimes 

apiculate. Styles 2- or 3-branched, unbranched portion flattened 

and fimbriate on margins (2-branched style) or slender, 3-angled 

basally, and glabrous (3-branched style), disarticulating from the 

summit of the achene. Achenes biconvex, lenticular or trigonous, 

obovate, elliptic-obovate or oblong, smooth, cancellate, reticulate, 

verrucose or warty.

A genus of ~300 species found in warm temperate and tropical climates worldwide, with the center of species diversity in 

southeastern Asia; ~15 species found in Venezuela.

Reference. Kral (1971).

Fimbristylis complanata (Retz.) Link, Hort. Berol. 1: 292. 1827; 

Kral, in Berry et al., Fl. Venez. Guayana 4: 572, fig. 453. 

1998. Scirpus complanatus Retz., Observ. Bot. 5: 14. 1791 

[1789]. Fimbristylis autumnalis var. complanata (Retz.) 

Kük., Bot. Jahrb. Syst. 59: 50. 1924.

FIGURE 8

Caespitose or rhizomatous perennial, 20–60(–100) cm tall; 

rhizomes short, ascending. Culms erect, compressed-trigonous 

proximally, flattened distally, firm, flexuous, sometimes twisted, 

antrorsely scabrid on margins (at least distally), 0.7–2 mm wide. 

Leaves numerous, subdistichous, ascending to spreading; sheaths 

short, uppermost dorsally keeled, brown to stramineous; ligule 

a horizontal band of dense hairs, 0.3–0.5 mm wide; blades linear, flattened, 5–70 cm × (1–)1.3–4 mm, adaxially antrorsely 

scabrous at base, smooth otherwise, margins finely and closely 

scabrous, abruptly narrowed to a flattened, slightly curved, acute 

tip. Inflorescence a terminal, compound, open or contracted 

anthela with ascending rays, 1.5–9 cm in diameter; involucral 

bracts 4–6, lowermost one leaflike, often exceeding the inflorescence, uppermost linear-setaceous; primary rays 4–12, unequal; spikelets 15–100 or more, ovoid-lanceoloid to lanceoloid, 

4–7(–9) × 1–1.5(–2) mm, acute, obtuse at base, solitary at ray 

tips; scales ovate to ovate-lanceolate, 1.4–2.5 × 1–1.5 mm, dorsally acute, brown, reddish-lineolate, midcosta 3-nerved, green, 

prolonged beyond the acute apex as a short, slightly excurved 

mucro. Stamens 2 or 3; anthers 0.7–1 mm long, apiculate. Styles 

3-branched, unbranched portion smooth, trigonous-dilated at 

base. Achenes trigonous with shallowly convex faces, obovoid, 

0.7–1 × 0.5–0.7 mm, apiculate, obtuse at base, tricostate, finely 

striolate-reticulate, tuberculate to nearly smooth, chalk-white 

when immature, gray-white or pale brown at maturity.

An often weedy species found in tropical and subtropical 

regions worldwide. In the Americas, known from Mexico, Central America, the West Indies, and South America (Colombia, 

Venezuela, the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, 

Argentina, and Uruguay). Widespread in Venezuela (Amazonas, 

Anzoátegui, Apure, Barinas, Bolívar, Delta Amacuro, Distrito 

Federal, Falcón, Lara, Mérida, Monagas, Nueva Esparta, Sucre, 

Táchira, Trujillo, and Zulia; Map 72). In the park, known from 

wet areas in montane forest on the south slope of Guaramacal; 

1,800–2,000 m.

MAP 72. Fimbristylis complanata occurrence in Venezuela.
NUMBER 100 • 6 9

FIGURE 8. Cyperaceae, Fimbristylis complanata (Retz.) Link. A. Habit. B. Inflorescence. C. Spikelet. D. Spikelet scale. E. Flower. F. Mature 

flower with developing achene. G. Achene. (A–G, Stergios et al. 20715).
7 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Oreobolus R. Br.

Oreobolus R. Br., Prodr. 235. 1810.

Schoenoides Seberg, Willdenowia 16: 181. 1986.

Perennial herbs, cushion- or mat-forming; rhizomes ascending to erect. Culms short, firm and rigid, obtusely angled or crescentform in cross section. Leaves primarily basal, distichous or 

indistinctly so, equitant; sheaths eligulate, open, strongly dilated, 

pale brown to purplish-brown, chartaceous; pseudopetioles 

linear-oblong, ± canaliculate adaxially; blades short, linear or 

trigonous, stiff and somewhat cartilaginous. Inflorescence terminal, racemose with several lateral spikelets or reduced to a single 

spikelet, short, generally hidden among the leaves and involucral 

bracts; involucral bracts leaflike; spikelets with 3 or 4 obscurely 

distichous scales, 1-flowered, terminal scale fertile, those below 

empty; scales deciduous from the rachilla, subcartilaginous, innermost one submembranous. Flowers bisexual, with 2 alternating whorls of scalelike tepals at base that are often persistent, 

rarely with bristles. Stamens 3. Styles 3-branched. Achenes obtusely trigonous, obovoid, ellipsoid or pyriform, surface smooth, 

glossy.

Found in Central America, South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Chile, including the Juan Fernandez Islands, and Argentina), the Falkland Islands, Malesia 

to southeastern Australia, Tasmania, New Zealand, Tahiti, and 

Hawaii; ~16 species, 3 of which are found in Venezuela.

References. Chacón et al. (2006); Seberg (1988).

Oreobolus is confined to high elevations in the tropics, and 

its disjunct distribution reflects a Gondwanian origin (Chacón et 

al., 2006).

Oreobolus venezuelensis Steyerm., Bol. Soc. Venez. Ci. Nat. 11: 

308, fig. 1. 1950.

FIGURE 9

Densely caespitose, 5–16 cm tall, often elevated on a simple or branching, slender woody caudex surrounded by fibrous 

roots and shrouded by old leaf sheath bases. Culms often hidden among the leaves, crescentform, scabrous on margins apically, often spirally twisted, glossy green, subtended by a large 

2-keeled prophyll at uppermost node. Leaves spirodistichous; 

sheaths 5–7-veined, reddish-brown; pseudopetiole 11–18 mm 

long; blades linear, crescentform, 20–46 × 0.8–1.4 mm, with 

(1–)2(–3) inconspicuous veins abaxially, often 2-veined medially adaxially, acute apically. Inflorescence a small racemose 

panicle of 2–5 spikelets from the upper leaflike bracts or sometimes reduced to a single spikelet; spikelets 3–6 × 1.1–1.5 mm; 

scales ovate-lanceolate, 3–5 × 1–1.5 mm, green-glaucescent, redlineolate, often tinged with purple, midcosta of the lowermost 

1 or 2 scales with prolonged foliar-like awns 3–14 mm long, 

innermost scale 2.7–3.5 × 0.9–1.4 mm, midcosta not prolonged 

beyond the acuminate apex or only shortly so as a sharp tip; 

tepals narrowly deltate, ovate or oblong-ovate, 0.9–1.3 × 0.3–

0.4 mm, ochroleucous. Anthers 1–2 mm long. Achenes obpyriform, 1.2–1.7 × 0.9–1.1(–1.3) mm, 3-sulcate and longitudinally 

wrinkled, truncate apically and bearing the blackened nub of the 

base of the style centrally, smooth, brown to reddish-brown, pale 

basally.

Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, found only in the Andes (Táchira and Trujillo; Map 73). Páramo de Guaramacal; 2,950 m.

Rhynchospora Vahl

Rhynchospora Vahl, Enum. Pl. 2: 229. 1805, nom. cons.

Perennial or sometimes annual herbs, vegetatively diverse; 

roots fibrous. Culms caespitose or borne singly, trigonous or 

obscurely so, occasionally cylindric, shallowly to deeply channeled along 1 side or margins (at least distally) with channel 

edges often antrorsely scabrous, essentially smooth, sometimes 

scabrous distally, glabrous or sometimes hirsute. Leaves basal 

or basal and cauline, rarely strictly cauline; sheaths green, light 

brown or reddish-brown, sometimes whitened at base, inner 

band membranous on basal sheaths, often purple-dotted, splitting with age, often membranous only at orifice on cauline 

sheaths, orifice truncate or concave, sometimes U-shaped, rarely 

convex; ligule absent or sometimes present, often a narrow band 

of thickened tissue or trichomes at adaxial junction of sheath 

and leaf blade; blades flattened, V-shaped or folded, occasionally 

involute or crescentform-capillary, linear, filiform or sometimes 

capillary, herbaceous or occasionally stiff, glabrous, hirsute, or 

occasionally scabrous or scabridulous distally, rarely papillose 

or transversely rugulose, margins and abaxial midvein usually 

antrorsely scabrous, ciliate or with setose hairs. Inflorescence terminal or both terminal and with a series of lateral partial panicles 

MAP 73. Oreobolus venezuelensis occurrence in Venezuela.
NUMBER 100 • 7 1

FIGURE 9. Cyperaceae, Oreobolus venezuelensis Steyerm. A, B. Habit. C. Inflorescence (detail). D. Spikelet. E. Spikelet scale. F. Flower.

G. Achene. (A–G, Niño et al. 1379.)
7 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

from the upper leaflike bracts, paniculate, corymbose, racemose 

or congested and head-like; involucral bracts in capitate species 

sometimes whitened at base; branches cylindric, 3-angled, crescentform or subcompressed, finely ribbed, scabrous, ciliate or 

smooth on margins, spikelets solitary or in fascicles at branch 

tips; spikelets ovoid, ellipsoid, lanceoloid or fusiform, cylindric or subcompressed, primarily 1- to several-flowered, scales 

spreading with maturing achenes; scales spirally imbricate, ovate 

to lanceolate, shallowly to deeply cymbiform, often inrolled 

around flower, finely and indistinctly nerved on sides, with a 

single distinct or sometimes indistinct wirelike midcosta, light to 

dark brown, or castaneous or ferrugineous, sometimes whitish. 

Flowers bisexual above the (1–)2–5(–9) empty basal scales of 

spikelet, terminal often staminate with a rudimentary ovary or 

reduced and empty; bristles (1–)6(–20), rudimentary or absent, 

when present antrorsely or retrorsely barbed, sometimes smooth 

or plumose. Stamens 1–3(–12); anthers linear, narrowly elliptic 

or oblong, often with minute crystalline papillae or lobes at base. 

Styles subulate, 2-branched or undivided, often long-exserted beyond apex of subtending scale. Achenes biconvex to subcylindric, sometimes inrolled with winged or wavy margins, obovate, 

oblong-obovate or oblong-elliptic, deeply pitted, transversely 

rugulose or sometimes smooth, expanded, usually triangular, or 

sometimes discoid, style base persistent at summit.

A subcosmopolitan genus of 250–300 species with its greatest species diversity in the Americas, especially warm temperate 

North America and in the neotropics. In the Americas, the genus 

is found in North America, Mexico, Central America, the West 

Indies, and South America (all country-level political units except Chile). Approximately 110 species are found in Venezuela.

References. Kükenthal (1949); Thomas et al. 

(2009).

KEY TO THE SPECIES OF RHYNCHOSPORA

1a. Inflorescence a terminal globose head of spikelets without lateral branching; involucral bracts scalelike, indurate, imbricately arranged, and forming an involucre around spikelets; achenes antrorsely scabrous at apex; bristles plumose . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. globosa

1b. Inflorescence a single spikelet or series of fascicles or panicles of spikelets, at least some branching evident; involucral 

bracts subtending each panicle or partial panicle 1(–2), leaflike, herbaceous; achenes smooth at apex; bristles antrorsely 

spinulose or wanting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Spikelet scales light-colored, pale, stramineous or light brown on sides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Leaf blades filiform, wiry, obtusely triangular-channeled or subcrescentform, 0.3–0.6 mm wide, often spirally 

twisted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. gollmeri

 3b. Leaf blades flattened or plicate, 1.3–30 mm wide, not spirally twisted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Leaves 5–7 in number, spaced at relatively long intervals, basal and cauline, usually all with well-developed 

blades; ligule or ligular line absent; blades 11–30 mm wide; spikelets 3–5(–5.5) mm long . . . . . R. locuples

 4b. Leaves >10 in number, spaced at relatively short intervals (at least at midculm), primarily cauline, basal 

leaves often with short blades or represented by sheaths only; sheaths ligulate with a thin band or line of 

tissue at adaxial junction of sheath and blade; blades 1.7–10.5 mm wide; spikelets 3.8–8 mm long . . . . .5

 5a. Style base triangular, 0.4–1 mm long; achenes obovate or rounded-obovate, 0.8–1.2 × 0.6–0.8 mm, with 

7–10 rows of horizontally oriented rectangular cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. polyphylla

 5b. Style base lanceolate, 1–2.3 mm long; achenes squarish-obovate, 1.1–1.6 × 0.8–1.2 mm, with 14–18 

rows of horizontally oriented rectangular cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. tuerckheimii

2b. Spikelet scales dark-colored, brown to dark brown or reddish-brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 6a. Achene surface transversely rugulose (wrinkled) with vertically oriented linear cells . . . . . . . . . . . . . R. rugosa

 6b. Achene surface cancellate or faintly so, with horizontally oriented rectangular cells in vertical rows . . . . . . . 7

 7a. Achene margins shallowly sulcate (channeled); sheaths and leaf blades often pilose . . . . . . . . . . R. lechleri

 7b. Achene margins rounded; sheaths and leaf blades glabrous or at most with some appressed pubescence . . . . 8

 8a. Basal and lower leaf blades on culm 9–23 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

 9a. Panicles open and very diffuse, terminal panicle 11–23 × 8–28 cm; scales light brown to brown; 

achenes with 15–19 vertical rows of horizontally oriented rectangular cells . . . . . . . . . R. immensa

 9b. Panicles dense, terminal panicle 3–13 × 3–8 cm; scales dark brown or reddish-brown; achenes with 

11–16 vertical rows of horizontally oriented rectangular cells . . . . . . . . . . . . . . . . . . . R. ruiziana

 8b. Basal and lower leaf blades on culm 2–8 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

 10a. Leaves >10 in number, spaced at relatively short intervals (at least at midculm), primarily cauline, 

basal leaves often with short blades or represented by sheaths only; sheaths ligulate with a band or 

thin line of tissue at adaxial junction of sheath and blade . . . . . . . . . . . . . . . . . . R. tuerckheimii

 10b. Leaves 6–11 in number, spaced at relatively long intervals, basal and cauline, usually all bearing 

well-developed blades; sheaths eligulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
NUMBER 100 • 7 3

 11a. Inflorescence partial panicles open, diffusely branched, pyramidal . . . . . . . . . . . . . . . . R. sp. A

 11b. Inflorescence partial panicles contracted, densely branched, narrowly ovoid or trullate . . . . 12

 12a. Spikelets 4–7 mm long; achenes obovate, 0.7–1.1 mm wide, obtuse to subtruncate at apex; 

style base sessile with achene apex, ends of base not upturned . . . . . . . . R. macrochaeta

 12b. Spikelets 6.5–9 mm long; achenes shortly oblong-obovate, 1.1–1.3 mm wide, truncate 

at apex with often slightly upturned shoulders; style base articulate with achene body, 

flaring basally, ends slightly upturned . . . . . . . . . . . . . . . . . . . . . R. guaramacalensis

Rhynchospora globosa (Kunth) Roem. & Schult., Syst. Veg. 2: 

89. 1817. Chaetospora globosa Kunth, in H. B. K., Nov. 

Gen. Sp. [fol.] 1: 185. 1815 [1816]; ibid. [qu.] 1: 230. 1815 

[1816].

Our material is referred to the nominate variety of the 

species.

Rhynchospora globosa var. globosa

Rhizomatous perennial, 30–115 cm tall; rhizomes short; 

culms solitary or borne tightly together in a row along rhizome 

or sometimes caespitose; roots coarse, 1–3 mm thick; sheathing 

base of culm 7–22 mm wide. Culms ascending, 0.7–2.3 mm wide, 

trigonous to obtusely trigonous, hardened, coarsely ribbed, green, 

glabrous, clavate at apex just below inflorescence head. Leaves 

6–30 per culm, primarily basal, often 1–3 cauline just above base, 

stiffly erect to ascending, 8.5–70 cm long; sheaths eligulate, short, 

stiff, coriaceous, indistinctly finely veined with indistinct cross 

partitions between veins, shiny, subsmooth and coriaceous abaxially, smooth with soft and spongy epidermis that is often crosspuckered adaxially, brown to brown-black or black, glabrous, 

inner band membranous, deeply U-shaped on upper sheaths, 

concave to truncate on lower sheaths; blades linear, 1–4 mm 

wide (flattened), with indistinct cross partitions between veins, 

V-shaped in cross section, subtrigonous distally, stiff, smooth, 

indistinctly finely veined, green, glabrous, margins remotely scabrous to smooth on older well-developed leaves, antrorsely scabrous to ciliate-scabrous on young leaves, ciliate-scabrous near 

base on cauline leaves. Inflorescence a single hemispherical to 

subglobose congested head of spikelets at summit of culm, 9–20 

× 8.5–22 mm; involucral bracts 5–9 (immediately surrounding 

spikelets, often more interspersed among spikelets), scalelike, 6–8 

× 2.8–6.5 mm, indurate, stiff and cartilaginous, broadly ovate to 

broadly rectangular or broadly obovate, apex obtuse to truncate 

or emarginate, light brown to yellow-brown, ciliate to glabrate 

along margins, lowest often leaflike with a blade 0.7–20 cm long, 

exceeding the spikelets; spikelets numerous, ovate-lanceolate to 

narrowly elliptic-lanceolate, often curvate, acuminate at apex, 

short-cuneate basally, 7–9 × 1.2–2 mm, densely aggregated; scales 

7–12 per spikelet, herbaceous to thickened and subcartilaginous, 

acute to narrowly obtuse in cross section, indistinctly very finely 

veined, uniformly light brown to brown or whitish stramineous, 

glabrous, margins scarious, acute to obtusely or acutely rounded 

at apex on lower fertile and sterile scales, acuminate at apex on 

upper fertile scales, midcosta pale green, indistinct, prolonged 

beyond apex as a short mucro on proximal scales, glabrous 

to ciliate-scabrous distally; fertile scales 4–6, ovate-elliptic to 

ovate-lanceolate, upper fertile scales narrowly ovate-lanceolate, 

(4.9–)5–8 × 1.5–4 mm; sterile scales 3–6 (excluding 2-keeled prophyll at base), ovate-lanceolate to ovate-elliptic, 2–4.5 × 0.8–3.4 

mm. Stamens 2 or 3; anthers 2.3–3.5 mm long, long-apiculate at 

apex, truncate and papillate basally. Styles entire to shortly bifid 

at tip, reddish. Achenes biconvex to subplano-convex, oblongobovate to narrowly oblong-obovate or obovate, (2–)2.1–2.8(–3) 

× 1.1–1.5 mm, minutely indistinctly cellular-reticulate (appearing 

smooth), light brown, cuneate to base, obtuse to subtruncate at 

apex, margins and sides at apex often with crystalline prickles; 

epidermal cells isodiametric to somewhat vertically elongated, 

no central nodules evident; style base triangular with 2 rounded 

basal lobes that overlap on either side of achene body, 0.6–1 × 

0.5–0.7 mm basally, pale brown, apex truncate with 2 teeth on 

each margin, sometimes with 2 medial smaller teeth as well; bristles 5–6, 3–5 mm long, equaling to overtopping style base (longer 

than achene body), plumose, with ascending hairs, reddish, apex 

antrorsely barbed.

Found in Mexico, Central America, the Greater Antilles 

(Cuba), and South America (Colombia, Venezuela, Trinidad and 

Tobago, the Guianas, Peru, Bolivia, Brazil, Paraguay, Argentina, 

and Uruguay). In Venezuela, found in Amazonas, Anzoátegui, 

MAP 74. Rhynchospora globosa var. globosa occurrence in 

Venezuela.
7 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Apure, Bolívar, Delta Amacuro, Guárico, Monagas, Portuguesa, 

and Trujillo (Map 74). In the park, collected on the south slope 

in an abandoned field near El Campamento below Cerro El Diablo; 1,800–2,000 m.

Rhynchospora gollmeri Boeckeler, Linnaea 37: 556. 1873.

Caespitose, glabrous perennial, 6–50(–65) cm tall, with fibrous remnants of old leaf sheath bases clothing base; rhizomes 

short, ascending. Culms erect, wirelike, often spirally twisted, 

subterete or obtusely trigonous, 0.4–0.8 mm wide, pale green. 

Leaves primarily basal, 1 or 2 lower cauline, 4–30 cm long; 

sheaths eligulate, short, lowermost often bladeless, brown, inner 

band submembranous, truncate or slightly concave at orifice, 

minutely red-dotted or short-lineolate (at least distally); blades 

filiform, wiry, obtusely triangular-channeled or subcrescentform, 

0.3–0.6 mm wide, often spirally twisted, abaxially finely veined, 

adaxially smooth, finely cellular-reticulate, green, margins (edges 

of channel) antrorsely scabrous, attenuate. Inflorescence a terminal corymb and 1 or 2 remote, lateral, small corymbs, each with 

(1–)2–8 short-pediceled spikelets, subtended by a leaflike, linearsetaceous involucral bract; spikelets ovoid-ellipsoid, 3.5–4.5 × 

1.1–1.5 mm, narrowly acute to short-acuminate; fertile scales 4 

or 5, ovate-elliptic to ovate-lanceolate, 2.5–4.2 × 1.4–2.3 mm, 

brown, blackish-lineolate, glabrous, margins narrowly scarious, mucronate at apex. Stamens 2 or 3; anthers 1.1–1.5 mm 

long. Styles 2-branched. Achenes biconvex, obovate or squarishobovate, 1.5–2 × 1–1.2 mm, subtruncate and shallowly constricted with thickened edges at apex, short-cuneate basally, 

finely and shallowly transversely rugulose, brown to red-brown; 

style base deltate to deltate-lanceolate, 0.6–0.7 × 0.5–0.8 mm 

basally, finely scabrous on edges proximally, pale brown; bristles 

5–6, very finely antrorsely barbed with short translucent hairs 

basally, shorter than to equaling apex of achene.

Found in South America (Venezuela and possibly Brazil). In 

Venezuela, found in Anzoátegui, Bolívar, Distrito Federal, Sucre, 

Táchira, and Trujillo (Map 75). In the park, known only from 

the area just below the Páramo de Guaramacal; 2,950 m.

Our collection differs from typical material in having short 

culms that are hidden among the leaves. However, characteristics of the leaves, inflorescence, flowers, and fruits do not otherwise significantly differ from typical material, and our collection 

probably represents only an ecological extreme of this species.

♦ Rhynchospora guaramacalensis M. T. Strong, Acta Bot. Venez. 

29: 208, fig. 1. 2006; Briceño, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 2: 471. 2010.

FIGURE 10

Rhizomatous perennial, 45–110 cm tall; rhizomes short, 

horizontal to ascending, woody. Culms triquetrous or shallowly 

so, with blunt angles, flexuous, green to dark green with pale 

ribs, smooth on angles. Leaves 3–7, basal and cauline, 16–70 cm 

long; sheaths eligulate, elongate, green or pale brown proximally, 

inner band membranous, reddish-brown, sparsely and finely 

appressed-pubescent to glabrous, orifice concave or subtruncate; 

blades narrowly linear, 2–4.5 mm wide, subplicate with obtuse 

or subrounded pleats, smooth on margins proximally, scabrous 

on abaxial midvein and margins distally, often sparsely setose 

adaxially on midvein, long-attenuate to triquetrous setiform tip. 

Inflorescence a terminal panicle and series of 2 or 3 lateral, remote, narrowly trullate, partial panicles from the upper leaflike 

bracts; terminal panicle 4–8 × 1–2 cm, lateral ones successively 

smaller toward the base, terminating elongate, very slender, 

MAP 75. Rhynchospora gollmeri occurrence in Venezuela. MAP 76. Rhynchospora guaramacalensis occurrence in Venezuela.
NUMBER 100 • 7 5

FIGURE 10. Cyperaceae, Rhynchospora guaramacalensis M. T. Strong. A. Habit. B. Inflorescence branch (detail). C. Spikelet. D. Spikelet scale.

E. Flower. F. Achene. (A, B, D, E, Stergios & Caracas 19742; C, F, Stergios & Caracas 19837.)
7 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

erect to ascending, flexuous peduncles; branches narrowly wingangled, pale green medially on sides, sparsely ascending-setose 

or smooth on angles; spikelets lanceoloid, 6.5–9 × 1–1.5 mm, 

3- or 4-flowered, acuminate at apex, cuneate basally; fertile 

scales lanceolate, 3.5–7 × 1.7–3 mm, dorsally obtuse to subrounded, submembranous, dark brown or dark reddish-brown, 

midcosta wirelike, greenish, prolonged beyond the acute to 

acuminate apex as a 0.5–2 mm long slightly recurved awn. Stamens 3; anthers 2.3–3 mm long. Styles shortly 2-branched at 

apex, long-exserted. Achenes thickly biconvex, shortly oblongobovate, 1.6–1.9 × 1.1–1.3 mm, truncate at apex with often 

slightly upturned shoulders, cancellate with 14–19 vertical rows 

of horizontally oriented rectangular cells, often semiglossy, pale 

brown to brown; style base narrowly triangular, articulate with 

achene body, attenuate, 1.3–1.7 × 0.6–0.8 mm, flaring basally, 

ends slightly upturned; bristles 3–5, equaling or slightly exceeding the achene body but not surpassing the tip of the style base, 

antrorsely scabrous, reddish.

Endemic to the Andes of Venezuela (Trujillo; Map 76). This 

species is, at present, known only from the park, where it has 

been collected on the south slope in Qda. Jirajara and in the 

Páramo de Guaramacal; (2,200–)2,800–3,100 m. 

This species differs from Rhynchospora macrochaeta in 

having longer spikelets, broader achenes with upturned shoulders, and a style base that is flared.

Rhynchospora immensa Kük., Bot. Jahrb. Syst. 56(Beibl. 125): 

17. 1921. Rhynchospora aristata var. immensa (Kük.) Kük., 

Bot. Jahrb. Syst. 74: 401. 1949.

Rhynchospora aristata auct., non Boeckler; Bono, Cat. Fl. Vert. Occid. Cordillera Mérida 92. 2010.

Rhynchospora hieronymi Boeckeler subsp. hieronymi auct., non Boeckeler; 

Dorr et al., Contr. U.S. Natl. Herb. 40: 45. 2000 [2001].

Robust, rhizomatous perennial, 115–220 cm tall; rhizomes 

short, stout, hardened; sheathing base of culm 1–3 cm wide. 

Culms ascending, 3.8–8 mm wide, narrowing to 2 mm distally, 

triquetrous, stiff and hardened, smooth, green to pale green distally, often golden brown proximally, glabrous or sometimes 

sparsely pubescent at nodes. Leaves 9–15, basal and cauline, 40–

110 cm long; sheaths eligulate, loose, light brown and semiglossy 

abaxially, golden-brown and highly glossy adaxially, glabrous, 

inner band herbaceous to thinly herbaceous on proximal sheaths 

with a deeply U-shaped orifice at apex; blades linear-lanceolate, 

9–23 mm wide, flattened to subplicate distally, green with the 

midvein often golden brown, closely and often indistinctly 

septate-nodulose, essentially glabrous, margins finely antrorsely 

scabrous, long-acuminate. Inflorescence a series of 4–6, open 

and very diffuse partial panicles from the upper leaflike bracts, 

terminal panicle 11–23 × 8–28 cm; involucral bracts leaflike, 

lowermost elongate, distal ones becoming successively shorter; 

primary inflorescence branches often finely antrorsely spinulose 

on margins; spikelets range from 30 (on lowermost panicles) to 

140 or more (on terminal panicles), ovoid-lanceoloid, obtuse to 

cuneate basally, acuminate apically, 7–9 × (0.9–)1.1–1.7 mm; 

fertile scales 3–7, ovate-lanceolate, 4.7–6.5 × 2.7–3.7 mm, obtuse to acute, light brown to brown, glabrous to very minutely 

puberulent at apex on distal scales of spikelet, short-awned 

or mucronate on distal scales of spikelet. Stamens 3; anthers 

2.5–3 mm long. Styles entire to shortly bifid. Achenes biconvex, subrounded-obovate or widely oblong-obovate, 1.5–1.8 × 

1–1.3 mm, cellular-cancellate, broadly rounded to subtruncate 

at apex, abruptly contracted at base to short stipe, with 15–19 

rows of horizontally oriented rectangular cells, pale brown to 

brown; style base triangular-lanceolate, 1.7–2.5 × 0.6–0.8 mm 

basally, flattened, brown; bristles 2–4, several often rudimentary, 

overtopping achene, shorter than to equaling tip of style base, 

antrorsely barbed, reddish.

Found in Central America and South America (Colombia, 

Venezuela, Trinidad and Tobago, Guyana, and Brazil). In Venezuela, known from Amazonas, Aragua, Bolívar, Miranda, Sucre, 

Trujillo, and Zulia (Map 77). In the park, found on both slopes 

of Guaramacal; 2,000–2,750 m.

Rhynchospora lechleri Steud. ex Boeckeler, Linnaea 37: 633. 

1873; Thomas, in Berry et al., Fl. Venez. Guayana 4: 623–

624. 1998.

Caespitose perennial, 18–40 cm tall; rhizomes short, knotty. 

Culms erect, trigonous or obtusely so, channeled along 1 side 

distally, flexuous, pilose or sparsely so. Leaves primarily basal, 

1–3 cauline, 5–30 cm long; sheaths short, eligulate, pilose, light 

brown, with a membranous inner band that is concave at orifice, orifice with a glabrous reddish-brown rim; blades linear, 

1.3–3.2 mm wide, plicate, pilose at least proximally, dull green 

abaxially, semiglossy adaxially, attenuate to a triquetrous apex. 

Inflorescence a terminal panicle and series of 2 or 3 lateral, 

MAP 77. Rhynchospora immensa occurrence in Venezuela.
NUMBER 100 • 7 7

sublax, pyramidal or narrowly pyramidal partial panicles from 

the upper leaflike bracts; terminal panicle 2–4 × 1.5–3 cm, with 

9–60 spikelets, lateral ones successively smaller toward the 

base, terminating elongate, ascending peduncles; branches ascending to divergent; spikelets ovoid-ellipsoid, 3.6–5.3 × 1.2–2 

mm, 3–4-flowered; fertile scales ovate-lanceolate, brown, often 

with castaneous sides, glabrous, with broad scarious margins, 

midcosta wirelike, prolonged beyond the obtuse apex as a bristly mucro. Stamens 3; anthers 1–1.5 mm long. Styles shortly 

2-branched, long-exserted. Achenes biconvex, squarish-obovate, 

1–1.3(–1.5) × 0.7–1.2 mm, truncate at apex, obtusely narrowed 

to base, margins shallowly sulcate with the edges of channel 

often red, cancellate, with 11–16 vertical rows of horizontally 

oriented rectangular cells, often glossy, reddish-brown or dark 

brown; style base narrowly triangular, flattened, attenuate, 0.7–

1.1 × 0.5–0.9 mm basally; bristles 2–4(–6), often caducous, several sometimes equaling or slightly exceeding the achene body, 

but not surpassing the tip of the style base, antrorsely scabrous, 

reddish.

Found in South America (Colombia, Venezuela Ecuador, Bolivia, and Peru). In Venezuela, known from Amazonas, Bolívar, 

Falcón, Mérida, Monagas, Táchira, and Trujillo (Map 78). In 

the park, known from a single collection from the Laguna de los 

Cedros; 2,000 m.

Rhynchospora locuples C. B. Clarke, Bot. Jahrb. Syst. 34(Beibl. 

78): 5. 1904; Thomas, in Berry et al., Fl. Venez. Guayana 4: 

624. 1998.

Coarse, rhizomatous perennial, 110–200 cm tall; rhizomes 

short, stout; roots very coarse; sheathing base of the culm 1.5–3 

cm wide. Culms solitary, triquetrous, 4–11 mm wide, very 

finely antrorsely appressed-setose or scabridulous. Leaves 5–7, 

basal and cauline, 70–100 cm long; sheaths eligulate, elongate, 

loose, sparsely pubescent like the culms, inner band herbaceous, thinly so on proximal bands, orifice U-shaped; blades 

flattened to subplicate, 11–30 mm wide, green on both sides, 

glabrous adaxially, scabrid apically, abaxial surface glabrous or 

the veins sometimes scabrid. Inflorescence a terminal panicle 

and series of (3–)4–8 remote or distally subcontiguous, dense, 

pyramidal partial panicles from the upper leaflike bracts; terminal panicle 10–20 × 6–10(–20) cm, lateral panicles successively smaller below; panicle branches pilose with crystalline 

trichomes; spikelets narrowly ellipsoid-lanceoloid, 3–5(–5.5) × 

0.7–1.3 mm, 1-flowered, solitary at branch tips or in fascicles 

of 2 or 3; fertile scale ovate-lanceolate, 2.5–4.5 × 1.3–2.2 mm, 

stramineous to light brown, dull, glabrous, mucronate at the 

acute apex. Stamens 3; anthers 1.5–2.2 mm long. Styles shortly 

2-branched. Achenes biconvex, obovate or oblong-obovate, 

1–1.7 × 0.8–1.1 mm, truncate apically, faintly cancellate with 

12–15 vertical rows of horizontally oriented rectangular cells, 

smooth and lustrous, pale; style base narrowly triangular, lanceolate or ligulate, slightly flared basally, 1–2 × 0.5–0.6 mm basally, longitudinally wrinkled, whitish or stramineous; bristles 

1–5, variable in length, several often rudimentary, 1 or 2 sometimes exceeding length of achene body, rarely exceeding tip of 

style base, antrorsely barbed, reddish.

Found in Central America and South America (Colombia, 

Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, this species 

has a disjunct distribution and is found in the Andes (Táchira 

and Trujillo) and in the Venezuelan Guayana (Bolívar; Map 79). 

In the park, found on the south slope of Guaramacal; 1,900–

2,900 m.

MAP 78. Rhynchospora lechleri occurrence in Venezuela. MAP 79. Rhynchospora locuples occurrence in Venezuela.
7 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

This coarse Rhynchospora species can be distinguished 

readily by its broad leaf blades and large, dense panicles of small, 

pale spikelets.

Rhynchospora macrochaeta Steud. ex Boeckeler, Linnaea 37: 

632. 1873; Thomas, in Berry et al., Fl. Venez. Guayana 4: 

624. 1998; Briceño, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 2: 472, fig. 18. 2010.

Rhizomatous perennial, 25–75(–100) cm tall; rhizomes 

short, woody, knotty. Culms trigonous or subtriquetrous with 

blunt angles, rigid and hardened, smooth, sometimes scabrid just 

below terminal panicle. Leaves 7–11, basal and cauline, 20–60 

cm long; sheaths eligulate, elongate, pale brown to brown, essentially glabrous, inner band membranous on proximal sheaths, 

herbaceous except at orifice on cauline sheaths, reddish-brown 

to dark brown, orifice truncate to concave or U-shaped, sparsely 

appressed-pubescent; blades linear, 3–7 mm wide, flattened to plicate distally with obtuse or rounded pleats, green on both sides, 

glabrous, attenuate. Inflorescence a terminal panicle and series 

of 2–4 lateral narrowly ovoid partial panicles from the upper 

leaflike bracts; terminal panicle sometimes broadly ovoid or 

subpyramidal, 3.5–6 × 1–7 cm, lateral ones successively smaller 

toward the base, terminating ascending to arching peduncles; 

branches short, ascending; spikelets narrowly ovoid-ellipsoid, 

4–7 × 1–1.3(–1.5) mm, 2- or 3-flowered; fertile scales narrowly 

ovate-elliptic to ovate-lanceolate or lanceolate, acute, 2.8–6.3(–

6.8) × 1.3–2.3 mm, dark brown or dark reddish-brown with 

blackish lineations, apex with a short antrorsely scabrous awn. 

Stamens 3; anthers 2.3–3.5 mm long. Styles shortly 2-branched. 

Achenes biconvex, obovate, narrowly obovate or narrowly 

elliptic-obovate, 1.2–1.7 × 0.7–1.1 mm, obtuse to subtruncate 

apically, cancellate with 12–19 vertical rows of horizontally 

oriented rectangular cells, often glossy, reddish-brown; style 

base narrowly triangular, attenuate, 0.9–2 × 0.5–0.7 mm basally; 

bristles 2–5(–6), shorter to longer than the achene body, several 

sometimes equaling or sometimes surpassing the tip of the style 

base, antrorsely scabrous, reddish.

Found in Mexico, Central America (Costa Rica), and South 

America (Colombia, Venezuela, Ecuador, Peru, and Bolivia). Venezuelan material is from the Andes (Lara, Mérida, Táchira, and 

Trujillo), the Sierra de Perijá (Zulia), and the Venezuelan Guayana (Amazonas and Bolívar; Map 80). Subpáramo and páramo 

near the Laguna del Pumar and in the Páramo de Guaramacal; 

(2,600–)2,800–3,100 m.

Jørgensen and León-Yánez (1999: 433) consider this name 

to be a synonym of Rhynchospora ruiziana Boeckeler. The two 

names were published simultaneously, and if the two species are 

combined, R. ruiziana is the name that must be adopted. However, study of type material of R. macrochaeta shows that our 

species differs significantly from R. ruiziana in vegetative, inflorescence, and floral characteristics, and the two are treated here 

as distinct species.

Rhynchospora polyphylla (Vahl) Vahl, Enum. Pl. 2: 230. 1805; 

Thomas, in Berry et al., Fl. Venez. Guayana 4: 626. 1998. 

Schoenus polyphyllus Vahl, Eclog. Amer. 2: 5. 1798.

Rhizomatous perennial, (45–)85–135 cm tall; rhizomes 

short, knotty, sometimes ascending; sheathing base of culm 

2.5–8 mm wide. Culms ascending or clambering over other vegetation, 2.3–5.3 mm wide, obtusely trigonous, stiff, and hardened, light brown proximally, green distally, essentially glabrous. 

Leaves ascending to spreading, numerous, primarily cauline, 

internodes short midculm and distally (leaves closely spaced), 

4–38(–58) cm long, lower cauline blades shorter than mid to 

upper cauline ones; sheaths somewhat loose, short midculm, 

elongate proximally, herbaceous, green distally, light brown 

proximally, short-pilose to glabrous, inner band membranous, 

red-brown and finely appressed-pubescent on proximal sheaths, 

membranous only at orifice on distal sheaths, orifice truncate, 

often tomentose; ligule a thin line or band of tissue at adaxial 

junction of sheath and blade; blades linear, 3.7–10.5 mm wide, 

flattened-plicate, herbaceous, green on both sides, sparsely pilose 

to glabrous abaxially, scabridulous adaxially (at least distally), 

attenuate. Inflorescence a series of 2–7, open to subcontracted, 

partial pyramidal panicles from the upper leaflike bracts; terminal panicle largest, 3–8 × 2–6 cm, with 48–310 spikelets, lower 

panicles successively smaller; inflorescence branches sharply angled, angles pilose; spikelets narrowly ovoid-ellipsoid to ovoidlanceoloid, 3.8–5.5 × 0.7–1.4 mm, narrowly acute to acuminate 

apically, short-cuneate basally; fertile scales 6–11, ovate-elliptic 

to widely ovate-lanceolate, acute to short-acuminate, 2.5–4 

× 1.3–3.2 mm, dull, uniformly whitened-stramineous to pale 

brown, reddish-lineolate (often faintly so), glabrous, margins 

narrowly scarious, mucronate apically. Stamens 3; anthers 1–2.3 

mm long. Styles entire. Achenes biconvex, obovate or roundedobovate, 0.8–1.2 × 0.6–0.8 mm, rounded apically, cancellate 

MAP 80. Rhynchospora macrochaeta occurrence in Venezuela.
NUMBER 100 • 7 9

with 7–10 rows of horizontally oriented rectangular cells, redbrown; style base triangular, attenuate to apex, slightly sagittate 

(shallowly 2-lobed) basally, 0.4–1 × 0.5–0.7 mm basally, light 

brown, dotted or flecked with dark brown; bristles 0–6, several 

often rudimentary, shorter than to equaling middle of achene 

body, antrorsely barbed, reddish.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Trinidad and Tobago, 

the Guianas, Ecuador, Peru, and Bolivia). Widespread in Venezuela (Amazonas, Anzoátegui, Aragua, Barinas, Bolívar, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, 

Monagas, Portuguesa, Táchira, Trujillo, and Yaracuy; Map 81). 

In the park, collected at several localities on the south slope of 

Guaramacal; (1,300–)1,800–2,000 m.

This species can be separated from Rhynchospora tuerckheimii, which has a similar habit, by its smaller obovate or 

rounded-obovate achenes, 0.8–1.2 × 0.6–0.8 mm, with 7–10 

rows of horizontally oriented rectangular cells, and triangular 

style base. Rhynchospora tuerckheimii has squarish-obovate 

achenes, 1.1–1.6 × 0.8–1.2 mm, with 14–18 rows of horizontally 

oriented rectangular cells, and a lanceolate style base.

Rhynchospora rugosa (Vahl) Gale, Rhodora 46: 275. 1944; 

Thomas, in Berry et al., Fl. Venez. Guayana 4: 627. 1998; 

Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 2: 474–475, fig. 21. 2010. Schoenus rugosus Vahl, 

Eclog. Amer. 2: 5. 1798.

Caespitose perennial, 27–135(–140) cm tall; rhizomes short, 

knotty. Culms erect, (0.8–)1.2–2.5(–3) mm wide, trigonous to 

obtusely trigonous, very firm, smooth, glabrous. Leaves 5–15, 

primarily basal, 2–5 cauline, 9–55 cm long; sheaths eligulate, 

short, smooth, green, glabrous, inner band membranous on 

basal blades, minutely red-speckled, with truncate orifice, upper 

and cauline sheaths membranous only distally and bearing a 

truncate to concave orifice; blades (1.3–)1.7–4.5(–5) mm wide, 

flattened to subplicate or folded, smooth, green, glabrous, longacuminate. Inflorescence a terminal panicle and series of 1 or 

2 lateral, remote cymose-paniculate partial panicles from the 

upper leaflike bracts; partial panicles 1–8(–11) × 0.8–2.7(–3.5) 

cm, lower on flexuous peduncles, 1–6 cm long; spikelets in fascicles at branch tips, ovoid-ellipsoid, acute, short-cuneate basally, 

3.5–4.7(–5) × 1.1–1.7 mm; fertile scales 3–5, ovate to oblongovate, acuminate, (2.3–)2.5–3.5 × 1.8–2.8 mm, brown to dark 

brown or brownish-black, glabrous, margins broadly scarious, 

often erose, apex with a 0.1–0.5 mm long mucro. Stamens 3; 

anthers 0.6–1.7 mm long. Styles 2-branched, branches twice as 

long as unbranched portion, exceeding the scales and curling. 

Achenes biconvex, obovate to elliptic-obovate, 1.6–1.9 × 1–1.3 

mm, truncate apically, transversely rugulose, light brown or 

yellowish-brown; style base triangular, 0.6–1 × 0.6–0.8(–1) mm 

basally, minutely antrorsely scabrous along margins, pale brown 

to dark brown; bristles 6, antrorsely barbed, equaling to exceeding apex of style base, often setose basally.

Found in Mexico, Central America, the Greater Antilles, 

and South America (Colombia, Venezuela, Trinidad and Tobago, 

the Guianas, Ecuador, including the Galapagos Islands, Peru, 

Bolivia, Brazil, Paraguay, Argentina, and Uruguay); adventive in 

Europe, Africa, Madagascar, Asia, Malesia, and the Pacific region. Widespread in Venezuela (Amazonas, Anzoátegui, Apure, 

Aragua, Bolívar, Cojedes, Distrito Federal, Falcón, Lara, Mérida, 

Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, 

and Trujillo; Map 82). Collected near La Cueva below Cerro El 

Diablo; 1,800 m.

MAP 82. Rhynchospora rugosa occurrence in Venezuela.

MAP 81. Rhynchospora polyphylla occurrence in Venezuela.
8 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Rhynchospora ruiziana Boeckeler, Linnaea 37: 641. 1873; 

Thomas, in Berry et al., Fl. Venez. Guayana 4: 626. 1998; 

Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 2: 475–476, fig. 22. 2010.

Rhynchospora macrochaeta auct., non Steud.; Ortega et al., BioLlania 5: 40. 

1987.

Coarse, rhizomatous perennials, 55–165 cm tall; rhizomes 

short, woody. Culms trigonous or obtusely so, rigid and hardened. Leaves basal and cauline, 45–110 cm long; sheaths eligulate, elongate, brown, inner band thinly herbaceous, orifice 

concave on basal sheaths, often convex or broadly rounded 

on cauline sheaths and tomentulose; blades linear, flattened or 

subplicate, 12–22 mm wide, glabrous, attenuate. Inflorescence 

a terminal panicle and series of 4–9 lateral, densely branched, 

pyramidal partial panicles from the upper leaflike bracts; terminal panicle 3–13 × 3–8 cm, lateral panicles successively smaller 

toward the base, terminating elongate, erect to ascending peduncles; branches short, ascending-setose on angles; spikelets 

narrowly ovoid to lanceoloid, 4–8 × 1–1.5 mm, 3–4-flowered, 

acute to acuminate apically; fertile scales ovate-lanceolate, acute 

to acuminate, 2.6–7 × 1.6–2.8 mm, dorsally obtuse, submembranous, often dark brown or reddish-brown, with a 0.5–1 mm 

long slightly recurved awn apically. Stamens 3; anthers 1.2–2.8 

mm long. Styles shortly 2-branched. Achenes lenticular, obovate 

to elliptic-obovate, 1.4–1.9 × 0.9–1.4 mm, broadly rounded to 

subtruncate apically, cancellate with 11–16 vertical rows of horizontally oriented rectangular cells, often glossy, pale brown to 

brown; style base narrowly triangular, attenuate, 0.8–2(–2.5) 

× 0.4–0.7 mm basally; bristles 1–5, often exceeding the achene 

body but not surpassing the tip of the style base, antrorsely scabrous, reddish.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Ecuador, Peru, and Bolivia). Venezuelan 

collections are from Bolívar, Distrito Federal, Táchira, and Trujillo (Map 83). In the park, collected in the Páramo de Guaramacal; 2,800–3,000 m.

Rhynchospora tuerckheimii C. B. Clarke ex Kük., Bot. Jahrb. 

Syst. 74: 402. 1949.

Rhizomatous perennial, (50–)94–210 cm tall; rhizomes 

short, knotty, sometimes ascending; sheathing base of culm 3–5 

mm wide. Culms ascending, 1.4–3.3 mm wide, obtusely trigonous to trigonous distally, stiff and hardened, tan proximally, 

green distally, thinly pilose-puberulent to glabrous proximally. 

Leaves ascending to spreading, numerous, primarily cauline, 

internodes short midculm and distally (leaves closely spaced), 

18–35 cm long, lower cauline blades shorter than the mid to 

upper cauline ones; sheaths somewhat loose, short midculm, 

elongate proximally, green distally, tan proximally, thinly 

pilose-puberulent or densely so on inner band, inner band narrowly membranous on basal sheaths, herbaceous and membranous only at orifice on distal sheaths, often reddish-brown and 

cross-puckered, orifice truncate to convex, minutely black- or 

red-dotted; ligule a thin line or band of tissue at adaxial junction of sheath and blade; blades narrowly linear, 1.7–5.7(–6.5) 

mm wide, subflattened-plicate, green on both sides, adaxial 

midvein pilose, abaxial surface sparsely pilose to glabrous, adaxially smooth, attenuate. Inflorescence a series of 3–7, open 

to subcontracted, partial pyramidal panicles from the upper 

leaflike bracts; terminal panicle largest, 3–7 × 2.3–5 cm, with 

30–70 spikelets, lower successively smaller; branches pilose on 

angles; spikelets narrowly ovoid-ellipsoid, (4–)4.5–7.5(–8) × 

MAP 83. Rhynchospora ruiziana occurrence in Venezuela. MAP 84. Rhynchospora tuerckheimii occurrence in Venezuela.
NUMBER 100 • 8 1

1.3–2.2 mm, acuminate apically, cuneate basally; fertile scales 

5–7, ovate-elliptic to widely ovate-lanceolate, 3–7 × 2–3.2 mm, 

acute to narrowly acute, dull, uniformly pale or stramineous, 

sometimes with sides washed in brown, glabrous, margins narrowly scarious, mucronate apically. Stamens 2; anthers 1.5–2.6 

mm long. Styles entire, exserted. Achenes biconvex, squarishobovate, 1.1–1.6 × 0.8–1.2 mm, obtuse to subtruncate apically, 

cancellate with 14–18 rows of horizontally oriented rectangular 

cells, pale brown to light brown; style base lanceolate, sagittate 

(2-lobed) basally, 1–2.3 × 0.5–0.8 mm basally, thin, brittle, pale 

brown, dotted or flecked with brown; bristles 1–3, several often 

rudimentary, shorter than to equaling apex of achene body, antrorsely barbed, reddish.

Found in Mexico, Central America, and South America 

(Venezuela and Guyana). In Venezuela, known only from the 

Cordillera de Mérida (Trujillo; Map 84). Found in montane 

or cloud forest in various localities in the park, including Casa 

Vicuyal, the Laguna de los Cedros, Páramo del Pumar, Páramo 

de Guaramacal, Qda. Honda, and El Campamento below Cerro 

El Diablo; 1,800–2,600(–3,000) m.

Spikelet scale color of this species is typically pale or stramineous. However, in some material from Guaramacal, the sides 

of the scales are washed in brown. This material does not differ 

otherwise from the material with pale or stramineous spikelet 

scales and can be readily separated from other Guaramacal species of Rhynchospora with brown scales by the numerous leaves 

with short internodes. See the discussion under R. polyphylla

above for differences between it and this species.

Rhynchospora sp. A

Rhizomatous perennial, 100–200 cm tall; rhizomes short, 

knotty, sometimes ascending; sheathing base of culm 8–15 mm 

wide. Culms solitary or 2 or 3 together, ascending, 1.8–4 mm 

wide, triquetrous with blunt angles or flattened-trigonous distally, stiff and hardened, scabridulous to smooth, pale brown 

or tan proximally, green distally. Leaves 6–9, basal and cauline, 55–85 cm long; sheaths eligulate, elongate, somewhat 

loose, glossy and somewhat spongy-thickened proximally, 

often with some cross walls (partitions) between the veins, 

green distally, tan proximally, glabrous, inner band membranous on basal sheaths, herbaceous and membranous only 

at orifice on distal sheaths, often reddish-brown, orifice concave, rimmed with brown; blades linear, 3.8–8 mm wide, 

subflattened-plicate proximally, plicate distally, herbaceous, 

green on both sides, semiglossy, abaxial surface glabrous, adaxial surface smooth or scabridulous distally, adaxial midvein 

smooth, very finely and closely antrorsely scabrous on margins 

and abaxial midvein (at least distally), attenuate. Inflorescence 

a terminal panicle and series of 3 or 4, open, remote, pyramidal, partial panicles on elongate ascending peduncles from the 

upper leaflike bracts; terminal panicle largest, 6–13 × 5–10 cm, 

with 20–100 spikelets, lower successively smaller; branches 

2–6 per panicle, subterete and glabrous proximally, sharply 

angled distally, angles antrorsely scabrous; spikelets narrowly 

lanceoloid, 5–8 × 1–1.5 mm, acuminate apically, cuneate basally; fertile scales 3–4, ovate-elliptic to ovate-lanceolate, 3.7–6 

× 1.4–2.3 mm, dorsally rounded to broadly rounded, acute to 

narrowly acute or obtuse, submembranous, semiglossy, brown 

to reddish-brown, glabrous, margins narrowly scarious, mucronate or short-awned apically. Stamens 3; anthers 1.3–1.8 

mm long. Styles shortly bifid apically, exserted. Achenes biconvex, narrowly oblong-obovate, 1.7–2.2 × 0.8–1.2 mm, obtuse 

to broadly rounded or subtruncate apically, glossy, cancellate 

with 14–18 rows of horizontally oriented rectangular cells, pale 

brown to light brown; style base lanceolate, 1.4–2.5 × 0.6–1 

mm at the shallowly sagittate base, thickened, pale brown; bristles 1–4, several often rudimentary, extending to 1⁄2 the length of 

achene body to equaling or exceeding its apex, not exceeding tip 

of style base, antrorsely barbed, reddish.

Evidently endemic to Venezuela and at present known only 

from a suite of collections made in Guaramacal (Trujillo state; 

Map 85). Fila de Agua Fría; 2,700–2,800 m.

This undescribed species belongs in Rhynchospora sect. 

Paniculatae Boeckler.

Scleria P. J. Bergius

Scleria P. J. Bergius, Kongl. Vetensk. Acad. Handl. 26: 142. 1765.

Perennial or sometimes annual herbs; rhizomes, when 

present, horizontal or short and nodose, hardened and knotty, 

sometimes tuberous. Culms erect, elongating and sprawling, 

or scandent, trigonous or triquetrous, harshly scabrous to 

smooth, glabrous or pubescent, green. Leaves well developed 

at the middle and upper nodes, basal ones essentially bladeless; 

sheaths 3-angled, closed at summit, distinctly veined, apex of 

MAP 85. Rhynchospora sp. A occurrence in Venezuela.
8 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

the inner band with a rounded, obtuse or triangular contraligule 

with distinct, straight or anastomosing veins, margins thickened 

or cartilaginous, sometimes with a short to elongate scarious 

appendage; ligule absent or sometimes present; blades linearelongate or sometimes lanceolate, flattened-plicate to plicate 

or somewhat inrolled along margins, 3-costate, herbaceous, 

weakly to harshly scabrous on margins and costae, glabrous 

or sometimes pubescent. Inflorescence paniculate or spikelike, terminal or terminal and a series of axillary partial panicles from the upper leaflike bracts; panicle branches 3-angled, 

sometimes narrowly winged, scabrous or smooth, pubescent or 

glabrous; panicle bractlets linear-lanceolate or setaceous, often 

ciliate or scabrous on margins; spikelets unisexual or bisexual 

and androgynous, sessile or on pedicels to 1 cm long; staminate spikelets lanceolate or narrowly oblong-ovate, cylindric 

or subcompressed, many-flowered; staminate scales numerous, 

basal 2 or 3 often 2-ranked and sterile, like the sterile pistillate 

spikelet scales, upper fertile ones spirally imbricate, narrowly 

ovate to lanceolate, membranous; pistillate spikelets ovoid to 

ovoid-lanceoloid or ellipsoid, subcompressed, often becoming 

obovoid after expanding to the width of the mature achene, cylindric to subcompressed, with a single terminal flower that is 

usually subtended by a much reduced, lanceolate fertile scale, 

hidden by the uppermost well-developed sterile scale; pistillate 

spikelet scales 2-ranked, uppermost 2 broadly naviculiform or 

cupuliform and spreading widely with the developing achene, 

carina 1–3-nerved, lateral nerves indistinct. Flowers unisexual; 

hypogynium (when present) borne at base of achene, sessile or 

stipitate, smooth or crustaceous, entire or 3-lobed, sometimes 

3–9-tuberculate near base, lobes entire or dissected, sometimes 

reflexed; cupula supporting the hypogynium and ovary shallow 

and dish-like, 3-lobed or obtusely trigonous with smooth margins, or deep and cuplike, enveloping the hypogynium and often 

ciliate on margin. Stamens 1–3; anthers often prickly appendaged apically. Styles capillary, 3-branched, unbranched portion 

glabrous. Achene body globose or ovoid to ellipsoid, rounded 

or obtusely trigonous, rarely trigonous or triquetrous, sometimes subconic, with straight to recurved apex, apiculate, bony 

or crustaceous, white or sometimes variegated with purple, glabrous or pilose, surface smooth, rugose, reticulate, trabeculate, 

papillate, verrucose or warty.

A genus of 200–225 species found primarily in warm temperate and tropical regions worldwide. In the Americas, found in 

North America, Mexico, Central America, the West Indies, and 

South America (all country-level political units except Chile). 

Approximately 42 species occur in Venezuela.

References. Camelbeke and Goetghebeur (2002); 

Core (1936, 1948).

KEY TO THE SPECIES OF SCLERIA

1a. Slender perennial, 0.5–0.6 m tall; inflorescence interruptedly fasciculate-spicate; spikelets in nodding fascicles; spikelet 

scales coarsely setose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. distans

1b. Coarse perennial, 2–3 m tall; inflorescence paniculate; spikelets separate on lateral branches; spikelet scales glabrous

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tropicalis

Scleria distans Poir., in Lamarck, Encycl. 7: 4. 1806; Camelbeke 

and Goetghebeur, in Berry et al., Fl. Venez. Guayana 4: 644–

645, fig. 508. 1998.

Slender perennial, 15–60 cm tall; rhizomes scaly, horizontally creeping, 2–4 mm thick. Culm erect, triquetrous, 0.8–2.7(–

3) mm wide, glabrous, angles smooth, wirelike. Leaves 2–8; 

sheaths subinflated apically, unwinged, hirsute to glabrescent, 

angles smooth or hirsute; ligule absent; contraligule obtuse or 

slightly concave, with hirsute margins, inner band often hirsute 

in a longitudinal line medially; blades flattened to subplicate, 

1–20 cm × 1.7–4(–5) mm, abaxial and adaxial surface glabrous, 

margins, midvein, and lateral veins often hirsute or sparsely so, 

subabruptly narrowed to an acute or acuminate, subflattened 

apex. Inflorescence terminal, interruptedly glomerate-spicate 

with no lateral branching, 8–22 cm × 7–10 mm, composed of 

4–10 sessile fascicles of spikelets; fascicles 4–8 mm in diameter, 

with 3–7(–9) spikelets, reflexed at maturity; bractlets ciliate basally, margins with dark hairs; spikelets bisexual (androgynous), 

ovoid-ellipsoid, 3.5–5 × 0.8–1.3 mm; staminate scales lanceolate, 

membranous; pistillate scale ovate to ovate-lanceolate, 2–2.5(–3) 

× 1–1.5 mm (excluding awn), sparsely setose on sides, pale green 

to dark purplish-brown, carina 1-nerved, setose, prolonged 

MAP 86. Scleria distans occurrence in Venezuela.
NUMBER 100 • 8 3

beyond the acute apex as a setose awn. Staminate flowers with 

2 stamens; anthers 1.7–2.2 mm long, with a lanceolate, bristly 

appendage at apex. Achene body subglobose to broadly obtusely 

trigonous in cross section, 1.4–1.8 × 1–1.3 mm, broadly obtuse 

to rounded apically, apiculate, with a brownish or blackish apiculum, porose, smooth, white or grayish, with a stipe-like, trigonous base; style base deciduous; hypogynium inconspicuous, 

3-lobed, with setaceous awl-shaped lobes, these often impressed 

into margins of base.

Found in North America (USA), Mexico, Central America, 

the West Indies, South America (Colombia, Venezuela, Trinidad 

and Tobago, the Guianas, Ecuador, Bolivia, Brazil, Paraguay, 

Argentina, and Uruguay), Africa, and Madagascar. Widely distributed in Venezuela (Amazonas, Anzoátegui, Apure, Aragua, 

Barinas, Bolívar, Distrito Federal, Falcón, Guárico, Lara, Mérida, 

Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, 

and Trujillo; Map 86). In the park, known from wet areas on the 

south slope of Guaramacal near El Campamento below Cerro El 

Diablo; 1,800–2,000 m.

♦ Scleria tropicalis M. T. Strong, Harvard Pap. Bot. 11: 199, fig. 

1. 2007.

Rhizomatous perennial, 2–3 m tall. Culms ascending, triquetrous, 2.5–7 mm wide, stiff and hardened, scabrid on angles, 

smooth proximally to scabrid on sides distally, finely ribbed, 

green proximally, often red or tinged with red on angles and 

ribs and at nodes distally. Leaves cauline, spaced at 5–15 cm 

intervals; sheaths eligulate, 5–10 cm long, broadly winged 

from the decurrent margins of the blade bases, wing margins 

finely and closely retrorsely spinulose, essentially smooth on 

sides, finely veined, often red-tinged; blades ascending, 30–70 

× 13–26 mm, plicate, 3-costate, margins and abaxial midcosta 

finely and closely retrorsely spinulose, decurrent on margins of 

sheath, long-attenuate to a subulate tip, green and often tinged 

with red; contraligule convex to rounded, with thickened, dark 

reddish or blackish hispid margins, veins of the inner band running straight to the margins, not anastomosing. Inflorescence a 

terminal rhombic panicle and a series of 0–3 lateral narrower 

and successively smaller partial panicles at intervals of 5–10 cm 

from the upper leaflike bracts; terminal panicle 2–3 × 2–2.5 cm; 

panicle branches stiff, triquetrous to wing-angled, often red or 

red-tinged medially, mostly very finely antrorsely spinulose with 

some retrorse barbs intermixed on margins; bracts subtending 

each partial panicle leaflike, becoming successively smaller distally; bracts subtending branches linear-setaceous, 2–7 cm long; 

pistillate spikelets generally lowest on each secondary or tertiary 

branch with 1–3 staminate ones above, 4–5.5 × 1.5–2 mm, narrowly ellipsoid-lanceoloid, cuneate basally, acuminate apically 

with a single terminal floret subtended by a linear reduced scale, 

sterile scales 3 or 4 proximally; staminate spikelets lanceoloid, 

short-cuneate basally, obtuse apically, 3–4 × 1.2 mm, with 3 sterile scales proximally and 2 to several linear membranous fertile 

scales distally, each subtending 3 stamens; sterile pistillate scales 

ovate to broadly so, thinly herbaceous, dark reddish-brown, 

midcosta slender, prolonged as a mucro beyond the cusp-like 

acute apex, lateral nerves indistinct; sterile staminate scales like 

the pistillate except somewhat narrower and shorter. Flowers 

unisexual. Stamens 3; anthers 1.5–2.3 mm long (including the 

black subulate appendage at apex). Styles 3-branched. Hypogynium indistinct, shallowly 3-lobed, connate with achene body, 

minutely dotted along edges. Achenes ovoid, obtusely trigonous 

to broadly so, 3.5–4.3 × 3–3.3 mm, abruptly narrowed to an 

apiculus apically, truncate basally, smooth, uniformly white or 

tinged with green.

Found in South America (Colombia and Venezuela). In Venezuela, known only from the Andes (Trujillo; Map 87). In the 

park, found in wet pasture and disturbed areas bordering montane forest on the south slope in El Campamento below Cerro El 

Diablo; 1,800–2,000 m.

Uncinia Pers.

Uncinia Pers., Syn. Pl. 2: 534. 1807.

Tufted, rhizomatous or rarely stoloniferous, monoecious, 

perennial herbs. Culms central, erect or ascending, sharply 

trigonous to subterete, striate, smooth proximally, often scabrid 

below inflorescence, base often surrounded by bladeless sheaths. 

Leaves basal, tristichous; sheaths ligulate; blades narrowly linear, flattened or involute, scabrous on margins and abaxial 

midvein. Inflorescence a terminal, simple, linear-cylindric, contiguous spike, androgynous with pistillate spikelets proximally 

and a short staminate spikelet distally; involucral bracts mostly 

small and scalelike or a few lower ones leaflike, deciduous or 

persistent; spikelets with spirally arranged scales; scales ovate to 

oblong; staminate spikelet with few to many densely arranged 

MAP 87. Scleria tropicalis occurrence in Venezuela.
8 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

scales, each axillating a flower; stamens (1) 2 or 3; pistillate 

spikelets 1-flowered; rachilla vestigial, elongated beyond the 

beak of the perigynia and bearing a hooked rudimentary spikelet 

apically; ovary enclosed in an obtusely trigonous, bottle-shaped, 

herbaceous sac (perigynium); style 3-branched, incrassate basally, exserted from the apex of the perigynia; bristles absent; 

style 3-branched, base not distinct, often thickened and persistent. Fruit a trigonous, oblong to ovate, beaked achene, surface 

smooth.

As presently construed, a genus of ~60 species found in 

Mexico, Central America, the Greater Antilles, South America 

(Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Chile, including the Juan Fernandez Islands, and Argentina), Malesia, 

Australia, New Zealand, and the Pacific Islands (including Hawaii). Three species occur in Venezuela.

References. Gehrke et al. (2010); Hamelin (1958); 

Starr and Ford (2009); Starr et al. (2004, 2008); Waterway and 

Starr (2007); Waterway et al. (2009).

Although molecular data (Starr et al., 2004, 2008; Waterway et al., 2009; Gehrke et al., 2010) resolve Uncinia as monophyletic, it is embedded in the very large genus Carex and is a 

major element in a core unispicate clade (Waterway and Starr, 

2007; Starr and Ford, 2009) that also includes Cymophyllus

and unispiculate species of Kobresia and Carex. Sister to this 

core unispicate clade is the Schoenoxiphium clade composed of 

Schoenoxiphium Nees and species of Carex. These two clades 

together compose one of the four major clades identified in 

Carex, contain representatives of all five genera recognized in 

the Cariceae, and include ~230 species found in temperate and 

montane habitats of the Northern and Southern Hemispheres.

Uncinia hamata (Sw.) Urb., Symb. Antill. 2: 169. 1900; Kearns 

and Goetghebeur, in Berry et al., Fl. Venez. Guayana 4: 661. 

1998; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 478, fig. 24. 2010. Carex hamata Sw., Prodr. 

18. 1788.

FIGURE 11

Caespitose, 20–70 cm tall; rhizomes short, ascending or 

horizontal. Culms erect, subtriquetrous to obtusely trigonous 

with blunt angles, 0.6–1.5 mm wide, smooth, glabrous. Leaves 

6–12 per culm, narrowly linear, primarily basal, 2 or 3 lower 

cauline, 8–50 cm long; sheaths elongate, glabrous, inner band 

membranous with a U-shaped finely ciliate orifice; ligule a thickened, finely ciliate, narrow band of tissue, ~0.3 mm wide, at adaxial junction of sheath and blade, merging from margins in an 

obtuse to acuminate configuration; blades 1.7–4 mm wide, flattened or subplicate, adaxially coarsely scabrous along mid and 

lateral veins (at least distally), abaxially smooth, finely veined 

with coarser pale green mid and lateral veins, dark green otherwise, long-attenuate to triquetrous apex. Inflorescence spike 

6–18 cm × 2.5–3.5 mm, staminate tip 1–3 cm long; spikelet 

rachilla exserted for 3–6 mm, strongly uncinate apically; pistillate scales oblong-obovate, 4–6 × 1.7–2.2 mm, obtusely to 

acutely narrowed to apex, widest above the middle, gradually 

narrowed to base, submembranous, greenish, glabrous or scabridulous apically dorsally, midcosta indistinct, sides finely and 

closely pale-nerved, these often merging proximally, ending short 

of the apex. Stamens 3; anthers ~1 mm long. Perigynia planoconvex or compressed-trigonous, 2-margined, oblanceolate, obtuse, gradually to subabruptly narrowed to a finely black-nerved 

stipitate base, 4.5–6.5 × 0.7–1.6 mm, tightly enveloping achene, 

finely nerved, often faintly so, scabridulous distally, smooth 

proximally, margins bristly ciliate; achenes plano-convex, oblong, 2.7–4 mm long, obtuse apically, truncate basally, brown, 

with 3 very fine, pale, marginal nerves, puncticulate.

Found in Mexico, Central America, the Greater Antilles, 

and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, and Argentina). In Venezuela, found in Amazonas, 

Anzoátegui, Aragua, Bolívar, Distrito Federal, Falcón, Lara, 

Mérida, Miranda, Sucre, Táchira, Trujillo, and Yaracuy (Map 

88). In the park, found in open areas on both slopes of Guaramacal; 1,900–2,600 m.

Dioscoreaceae

L. J. DORR AND B. STERGIOS

Perennial geophytes, twining-climbing vines (our species), 

herbs or rarely shrubs; rhizomes short and often subtending 1 

to several tubers; aerial shoots often long. Leaves alternate (our 

species), opposite or whorled; leaf blades simple (our species) or 

digitately compound, usually cordate; margins entire, sometimes 

palmately lobed; principal veins arcuate, ascending from base to 

MAP 88. Uncinia hamata occurrence in Venezuela.
NUMBER 100 • 8 5

FIGURE 11. Cyperaceae, Uncinia hamata (Sw.) Urb. A. Habit. B. Inflorescence (detail). C. Flower. D. Spikelet scale. E. Achene. (A, Stergios 

et al. 19320; B–E, Stergios et al. 20634.)
8 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

apex, secondary veins reticulate; tendrils absent; petioles usually 

pulvinate at the base and apex; lateral nodal flanges (“stipules”) 

sometimes present at base. Inflorescences usually axillary, in panicles, cymes, spikes or racemes. Flowers 3-merous, epigynous, 

actinomorphic, unisexual or bisexual (plants dioecious, rarely 

monoecious). Tepals 6 in 2 whorls, ± free or connate at the base. 

Stamens (or staminodes) 6 in 2 whorls, the inner whorl sometimes sterile or absent; filaments free, rarely short-connate at the 

base; anthers 2-thecate, 4-sporangiate, longitudinally dehiscent. 

Gynoecium syncarpous, 3-carpellate; septal nectaries sometimes 

present; ovary (1–)3-locular, inferior; ovules anatropous, 2 to 

many per locule; placentation axile or parietal; nectaries septal 

or at the base of tepals; styles 3-branched or -lobed. Fruit a capsule (our species), rarely a samara or berry; seeds flattened and 

winged or globose and unwinged.

A principally tropical family of 2 genera and ~800(or 350–

400?) species found in North America (USA), Mexico, Central 

America, the West Indies, South America (all country-level political units), Europe, Africa, Madagascar, Asia, Australia, and the 

Pacific Islands. Dioscorea L. is the only genus of Dioscoreaceae 

found in Venezuela.

References. Al-Shehbaz and Schubert (1989); Caddick et al. (2002a, 2002b); Huber (1998); Knuth (1924); Merckx 

et al. (2010); Wilkin et al. (2005).

In Guaramacal, Dioscoreaceae and Burmanniaceae are the 

only families included in the Dioscoreales, and as discussed under 

Burmanniaceae, their circumscription has been problematic, 

with respect to which genera to include in each family and the 

limits of these genera. Caddick et al. (2002b), utilizing molecular 

data, argue that a monophyletic Dioscoreaceae can be defined 

by including four genera: Dioscorea s.l. (including Tamus L. and 

Rajania L.), Stenomeris Planch., Tacca J. R. Forst. & G. Forst., 

and Trichopus Gaertn. (synonym: Avetra H. Perrier). Stenomeris

and Tacca often are placed in their own monogeneric families, 

Stenomeridaceae and Taccaceae, respectively, whereas Trichopus and Avetra are segregated as Trichopodaceae. Merckx et 

al. (2010), however, argue that Dioscoreaceae sensu Caddick et 

al. (2002b) is not monophyletic, and they found strong support 

for Taccaceae and Trichopodaceae. We have chosen to adopt 

the circumscription of Merckx et al. (2010) and restrict Dioscoreaceae to Dioscorea and Stenomeris.

Dioscoreaceae have economic value as they are the source of 

edible yams, notably Dioscorea alata L., and medicinal (corticosteroid) and contraceptive compounds.

Dioscorea L.

Dioscorea L., Sp. Pl. 1032. 1753, nom. cons.

Lianas, herbaceous or suffruticose; aerial stems annual, 

creeping or climbing, twisting clockwise (dextrorse) or counterclockwise (sinistrorse; direction species specific); some species 

producing bulbils (aerial tubers) in the leaf axils; glabrous or 

pubescent with simple to stellate or T-shaped hairs. Leaves alternate (our species), opposite or whorled, simple (our species) or 

compound, entire (our species) or palmately lobed. Plants dioecious, rarely monoecious or with vestigial staminate or pistillate 

flowers. Inflorescences simple or compound, partial inflorescences usually pendent, spicate or racemose; staminate flowers 

solitary or paired, sometimes in cymules; pistillate flowers usually solitary or paired. Spicate inflorescences lax to dense, flowers usually with 2 floral bracteoles, usually unequal; tepals 6, in 

2 whorls, equal or unequal, free to basally connate, inserted on a 

disk of variable size and shape. Staminate flowers: stamens 3 or 

6, inner or outer whorl sometimes staminodial; pistillode present 

or not. Pistillate flowers: staminodes 3 or 6 or wanting; ovary 

inferior, 3-locular; ovules 2 per locule; styles 3, variously fused at 

base and free toward the apical stigmatic surfaces. Fruit a capsule 

(our species), 3-winged, loculicidally dehiscent, rarely baccate 

or a 1-winged samara. Seeds usually 6, 2 per locule, sometimes 

fewer; usually winged (our species) or sometimes wingless.

A pantropical genus of 350–400(or ~800?) species; some 

species also found in temperate parts of North America, Europe, 

and Asia. Approximately 40 species occur in Venezuela.

Knuth (1924) and Huber (1998), both relying on morphological characters, proposed scores of subgeneric groupings of 

Dioscorea, but whether or not these should be maintained is the 

subject of ongoing molecular and morphological research (Caddick et al., 2002b).

KEY TO THE SPECIES OF DIOSCOREA

1a. Staminate flowers sessile to short-pedicellate, solitary or in minute cymules; leaf bases cordate to ± hippocrepiform, sinuses 

broad; hairs, where present, simple and multicellular; stems twisting counterclockwise (sinistrorse) . . . . . . . D. meridensis

1b. Staminate flowers pedicellate, always solitary, never in cymules; leaf bases truncate to cordate, not hippocrepiform or 

with broad sinuses; glabrous or pubescent (hairs not multicellular); stems twisting clockwise (dextrorse) . . . . . . . . . . 2

2a. Leaf blades 9–11-nerved, glabrous above, pubescent (rarely glabrous) below on nerves; petioles >4 cm long . . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. lehmannii

2b. Leaf blades 5–7(–9)-nerved, glabrous above and below; petioles <3 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Stems terete; leaf blades 6–11 × 2.5–5 cm, 5–7-nerved; staminate flowers: pedicels 5–8 mm long; filaments <0.5 

mm long, separate at anthesis; staminal disk conspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. lisae

 3b. Stems angular to slightly winged; leaf blades 11–18 × 5–10.5 cm, 7–9-nerved; staminate flowers: pedicels 2–4 

mm long; filaments ~1 mm long, connate below, free above; staminal disk inconspicuous . . . . . . . D. coriacea
NUMBER 100 • 8 7

Dioscorea coriacea Humb. & Bonpl. ex Willd., Sp. Pl. 4(2): 794. 

1806.

Helmia moritziana Kunth, Enum. Pl. 5: 422. 1850. Dioscorea moritziana

(Kunth) R. Knuth, Notizbl. Bot. Gart. Berlin-Dahlem 7: 197. 1917.

Dioscorea sp. B; Dorr et al., Contr. U.S. Natl. Herb. 40: 46. 2000 [2001].

Suffruticose twining vines; stems to 3 mm in diameter, angular to slightly winged (at least when dry), twisting clockwise 

(dextrorse), minutely papillate. Leaf blades ovate-lanceolate, 

11–18 × 5–10.5 cm, base subcordate to truncate, apex acute 

to acuminate, 7(–9)-nerved, glabrous above and below, coriaceous; petioles 2.5–3.5 cm long, minutely papillate. Staminate 

inflorescences racemose, 1–4 racemes per node, to 15 cm long; 

rachis minutely pubescent to glabrescent; flowers solitary, shortpedicellate; pedicels 1.5–4 mm long. Staminate flowers: green, 

infundibuliform, tepals ± equal, reflexed at anthesis; stamens 3; 

staminal disk inconspicuous; filaments to 1 mm long, connate 

below, free above; staminodes 0; stamens concealing a pistillode. 

Pistillate inflorescences spicate, to 15 cm long (to 40 cm long 

in fruit); flowers solitary. Pistillate flowers: tepals ± equal, free, 

reflexed at anthesis; style fused below, separate above, stigmas 

± digitate; staminodes 0. Capsules oblong, 1.5–2 × 0.6–1.2 cm, 

base rounded to truncate, apex acute, glabrous.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, reported from Apure, Aragua, Barinas, 

Distrito Federal, Lara, Mérida, Portuguesa, Táchira, Trujillo, 

Yaracuy, and Zulia (Map 89). In the park, found in montane forest on both slopes of Guaramacal; 1,800–2,100 m.

Common Name. Ñame de monte.

Staminate inflorescences in the few Guaramacal collections 

we have of Dioscorea coriacea all have a single raceme per node, 

whereas outside the range of our flora staminate inflorescences 

have up to four racemes per node.

Dioscorea lehmannii Uline, Bot. Jahrb. Syst. 22: 430. 1897; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 

2: 487. 2010.

Suffruticose twining vines; stems to 4 mm in diameter, somewhat angled to very slightly winged (at least when dry), twisting 

clockwise (dextrorse), minutely pubescent. Leaf blades ovate to 

broadly ovate, 4–16 × 6–14 cm, base cordate, apex acuminate 

to cuspidate, 9–11-nerved, glabrous above, pubescent below on 

nerves (rarely glabrous), coriaceous; petioles 4.5–6.5 cm long, 

minutely pubescent to glabrescent. Staminate inflorescences paniculate, 1–8 racemes per node, 8–25 cm long; rachis minutely 

pubescent; flowers solitary, short-pedicellate; pedicels 2–4 mm 

long. Staminate flowers: green (or pink), campanulate to subrotate at anthesis, tepals equal; stamens 3; staminal disk prominent, annular; filaments short, erect, aggregated in the center of 

the disk, connivent; staminodes 0. Pistillate inflorescences spicate, to 12 cm long (to 30 cm long in fruit); flowers solitary. 

Pistillate flowers: tepals equal, free, spreading; style fused below, 

diverging toward apex, stigmas digitate; staminodes 0. Capsules 

ovate to narrowly ovate, 2–2.2 × 0.7–1 cm, base rounded, apex 

acute to acuminate, glabrous.

Found in South America (Colombia, Venezuela, and Ecuador). In Venezuela, known only from the Cordillera de Mérida 

MAP 89. Dioscorea coriacea occurrence in Venezuela. MAP 90. Dioscorea lehmannii occurrence in Venezuela.
8 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

(Portuguesa and Trujillo; Map 90). In the park, found on both 

slopes of Guaramacal; 1,900–3,100 m.

Knuth (1924) allied Dioscorea lehmannii with D. coriacea; 

the former is recognized by the presence of a staminal disk and 

the latter by its absence. He also placed these two species and D. 

moritziana (=D. coriacea) in D. sect. Cylcadenium Uline.

♦ Dioscorea lisae Dorr & Stergios, Sida 20: 1007, figs. 1, 2. 

2003; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 487–488, fig. 1. 2010.

FIGURE 12

Dioscorea sp. A; Dorr et al., Contr. U.S. Natl. Herb. 40: 46. 2000 [2001].

Herbaceous twining vines; stems delicate, ~1 mm in diameter, terete, twisting clockwise (dextrorse), glabrous. Leaf blades 

ovate-lanceolate, 6–11 × 2.5–5 cm, base cordate to truncate, 

apex acuminate to long-acuminate, 5–7-nerved, glabrous above 

and below, membranous; petioles 1.5–3.5 cm long, glabrous. Staminate inflorescences paniculate, with 1–4 racemose branches, 

8–13 cm long; rachis minutely papillate; flowers solitary, pedicellate; pedicels 5–8 mm long. Staminate flowers: greenish-red 

to purple, rotate at anthesis, tepals equal, free, slightly reflexed 

at anthesis; stamens 3; staminal disk conspicuous flattened, annular; filaments very short to ~0.25 mm long; staminodes 0. 

Pistillate inflorescences spicate, to 14 cm long; flowers solitary. 

Pistillate flowers: tepals equal; styles connate below, free above 

toward stigmatic surface; staminodes 0. Capsules elliptic, 2–3 × 

1.2–1.5 cm, apex and base rounded, glabrous.

Endemic to the Andes of Venezuela (Trujillo; Map 91). In 

the park, known only from montane forest on the north slope of 

Guaramacal; 1,800–3,000 m.

Several specimens of Dioscorea lisae have both infructescences and staminate inflorescences, indicating that this species 

is not strictly dioecious.

Dioscorea meridensis Kunth, Enum. Pl. 5: 334. 1850.

Suffruticose twining vines; stems to 5 mm in diameter, ± 

terete, grooved, twisting counterclockwise (sinistrorse), sparingly pubescent with simple, multicellular hairs, becoming 

glabrescent. Leaf blades orbicular to broadly ovate, 12–19 × 

8.5–16.5 cm, base cordate to ± hippocrepiform, sinus broad, 

3–4 × 2 cm, apex acute to cuspidate, 9–11(–13)-nerved, glabrous above, sparingly pubescent below with simple, multicellular hairs especially along nerves, membranous; petioles 

8–12 cm long, sparingly pubescent with simple, multicellular 

hairs. Staminate inflorescences paniculate, with 4–6(or more) 

branches, to 30 cm long; rachis pubescent with simple, multicellular hairs; flowers in minute cymules or solitary, sessile 

to short-pedicellate; pedicels ~1 mm long. Staminate flowers: 

green, short-campanulate at anthesis, tepals ± equal; stamens 

3; staminal disk inconspicuous; filaments short, distinct; staminodes 3; stamens and staminodes surrounding a pistillode. 

Pistillate inflorescences spicate, to 20(–40) cm long; flowers 

solitary. Pistillate flowers: not seen. Capsules ovate to ovateoblong, 1.8–2.2 × 1.3–1.5 cm, base cuneate, apex truncate, 

glabrous.

Found in South America (Colombia, Venezuela, Ecuador, and Peru). In Venezuela, known from the Andes (Mérida, 

Táchira, and Trujillo) and the Sierra de Perijá (Zulia; Map 92). 

Understory of montane forest on the north slope of the park; 

1,800–2,500 m.

MAP 91. Dioscorea lisae occurrence in Venezuela. MAP 92. Dioscorea meridensis occurrence in Venezuela.
NUMBER 100 • 8 9

FIGURE 12. Dioscoreaceae, Dioscorea lisae Dorr & Stergios. A. Habit (of a plant with mixed inflorescences, either male inflorescences or male inflorescences arising from the axils of infructescences). 

B. Staminate flower. C. Pistillate flower. (A, Dorr et al. 8921; B, C, Dorr & Barnett 8057.)
9 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Eriocaulaceae

L. J. DORR AND S. MIGUEL NIÑO

Perennial or rarely annual herbs, terrestrial (favoring very 

wet sites) or aquatic, rosulate, acaulescent or rarely caulescent. 

Rhizomes present or absent. Leaves alternate, spiral, rarely distichous or whorled, almost always basal, lanceolate to linear, flat 

or fleshy. Inflorescences in terminal heads, solitary or in umbels; 

involucral bracts usually present; peduncles long or short, enclosed at base by a tubular sheath. Flowers small, actinomorphic 

or zygomorphic, sessile or very short-pedicellate, numerous (10–

1,000 per head), unisexual or very rarely bisexual (Rondonanthus Herzog), some species with flowers of both sexes on the 

same head or in separate heads, rarely dioecious; hypogynous. 

Sepals 2 or 3, free or connate basally, sometimes completely connate. Petals 2 or 3 or absent, free or connate, eglandular or with a 

gland (Eriocaulon L.). Staminate flowers: stamens usually 3 or 2, 

or 6 or 4 in 2 whorls; filaments free or adnate to the corolla tube, 

sometimes with an androphore; anthers dorsifixed or basifixed, 

1- or 2-thecate, 2- or 4-sporangiate; pistillodes present. Pistillate 

flowers: ovary superior, 2- or more commonly 3-locular; ovules 

1 per locule, pendulous, orthotropous; styles 2 or 3 fused, stigmatic portions fused or not, style-like appendages nectariferous 

or not; staminodes present or absent. Fruit a capsule, generally 

loculicidally dehiscent, rarely an indehiscent achene. Seeds 1 per 

locule, rarely reduced to 1 per fruit; seed coat reticulate, striate, 

rough or smooth; endosperm copious.

A cosmopolitan family of 7 or 8 genera and ~1,200 species, 

with its greatest diversity in subtropical and tropical America, 

but also found in Europe, Africa, Madagascar, Asia, Australia, 

and the Pacific Islands. The majority of genera and species of 

Eriocaulaceae occur in mountainous areas of the Venezuelan 

Guayana and in west central and southeastern Brazil. In Venezuela as a whole, 6 or 7 genera and ~100 species occur.

References. Andrade et al. (2010); Costa and Sano 

(2013); Giulietti et al. (2012); Kral (1989); Parra et al. (2010); 

Stützel (1998); Unwin (2004).

Paepalanthus Mart.

Paepalanthus Mart., Ann. Sci. Nat., Bot., sér. 2, 2: 28. 1834, nom. cons.

Tonina Aubl., Hist. Pl. Guiane 2: 856. 1775.

Lachnocaulon Kunth, Enum. Pl. 3: 497. 1841.

Paepalanthus subgen. Actinocephalus Körn., in Martius, Fl. Bras. 3(1): 321. 

1863. Actinocephalus (Körn.) Sano, Taxon 53: 99. 2004.

Blastocaulon Ruhland, in Engler, Pflanzenr. 4, 30 (Heft 13): 223. 1903.

Rosulate herbs, acaulescent to caulescent, erect or creeping; 

usually vegetative parts pubescent, trichomes simple or malpighiaceous (“T”-shaped). Leaves linear, membranous or coriaceous. Inflorescences a single head (pedunculate or sessile) or a 

group of heads terminating a rosette, heads in umbels or not 

(our species); involucral bracts present. Flowers unisexual and 

bisexual (plants polygamous), 2- or 3-merous, subtended by a 

single bract, usually resembling the sepals, perianth biseriate and 

involute. Staminate flowers: sepals free or connate basally, forming a membranous tube, glabrous or pilose; petals fused; stamens 

2-thecate (rarely 1-thecate), 4-sporangiate (rarely 2-sporangiate); 

filaments free; gynoecium reduced to glands. Pistillate flowers: 

sepals free or connate basally, usually pubescent; petals free (or 

absent), inserted ± at the same height as the stigmas and opposite 

to them, apex papillate; gynoecium with simple or bifid stigmas 

and appendages (glands).

A predominately neotropical genus of ~450 species found in 

North America (USA), Mexico, Central America, the West Indies, 

South America (Colombia, Venezuela, the Guianas, Ecuador, 

Peru, Bolivia, Brazil, and Paraguay), Africa, and Madagascar. 

Approximately 50 species occur in Venezuela, the majority in the 

Venezuelan Guayana.

References. Rosa and Scatena (2007); Ruhland 

(1903); Trovó et al. (2013).

Stützel (1998) argues that circumscription of Paepalanthus

is problematic as he interprets the genus as “a conglomerate of 

several, probably not closely related groups of species.” Andrade 

et al. (2010), however, identify a monophyletic Paepalanthus s.l. 

if Actinocephalus (Körn.) Sano, Blastocaulon Ruhland, Lachnocaulon Kunth, and Tonina Aubl. are considered synonyms 

(as is done here, but see below). Trovó et al. (2013) resolve the 

Paepalanthoideae as monophyletic but hint at a dismemberment 

of Paepalanthus s.l., which Costa and Sano (2013) have initiated by recognizing Actinocephalus as a genus endemic to Brazil. 

The subgeneric groupings of Paepalanthus s.l. proposed by Ruhland (1903) based on morphology are clearly artificial and do 

not agree with the relationships that now are being revealed by 

analyses of molecular data.

The generic synonymy proposed above creates a nomenclatural problem in that Tonina is a much older name than Paepalanthus, which is conserved against Dupatya Vell. and not Tonina. 

The ICN (McNeill et al., 2012: Art. 11) requires us to adopt the 

older of the two generic names, but adopting the Aublet name 

for the ~450 species of Paepalanthus would be destabilizing nomenclaturally. Fortunately, the ICN (McNeill et al., 2012: Art. 

56.1) also permits names to be rejected, and we expect that once 

the status of Paepalanthus is fully resolved a proposal to reject 

the monotypic Tonina will be made.

Paepalanthus pilosus (Kunth) Kunth, Enum. Pl. 3: 518. 1841; 

Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 2: 498–499, foto 8. 2010. Eriocaulon pilosum

Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 200. 1815 [1816]; 

ibid. [qu.] 1: 251. 1815 [1816].

FIGURES 13, 25D

Paepalanthus karstenii Ruhland, in Engler, Pflanzenr. IV, 30 (Heft 13): 155. 

1903; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 

494–495, fig. 2, foto 5. 2010.

Paepalanthus karstenii var. corei Moldenke, Phytologia 29: 386. 1975.
NUMBER 100 • 9 1

FIGURE 13. Eriocaulaceae, Paepalanthus pilosus (Kunth) Kunth. A. Habit. B. Inflorescence with peduncle and involucral bracts. C, D. Staminate flowers. C. Staminate flower with pistillodes barely equaling petal tube in length (anthers mostly removed, one filament visible). D. Staminate flower with three anthers exceeding petal tube in length. E, F. Pistillate flowers. E. Flower with two-locular ovary. F. Flower with 

three-locular ovary. (A–C, E, F, Dorr et al. 7324; D, Stergios et al. 18726.)
9 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Rosulate herbs, 1.5–2.5 cm tall, cushion-like or forming 

small colonies. Leaves recurved, 10–14(–20) × 1–2 mm, apex 

sharply acute, margins with scattered long cilia (or with scars 

where these hairs have fallen), otherwise glabrous. Inflorescences 

several, arising from lower nodes; peduncles 0.5–4(–9) cm long, 

apex pubescent, subtended by a 3-lobed, glabrous, sheathing 

bract; heads 2–5 mm in diameter. Involucral bracts light brown, 

apex pubescent, sometimes margins and abaxial surface also pubescent. Sepals black. Petals of the pistillate flowers oblanceolate, pilose on both surfaces but surfaces not completely covered 

by hairs.

Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, found only in the Andes (Lara, Mérida, 

Táchira, and Trujillo; Map 93). In the park, found in waterlogged soils, often with Sphagnum L. and other mosses, in the 

Páramo de Guaramacal and near the Laguna del Pumar; ~2,600–

3,100 m.

A critical revision of the species of Paepalanthus that occur 

in the northern Andes is needed. The last comprehensive treatment to deal with these species was that of Ruhland (1903), 

but a number of northern Andean species have been described 

since then without consideration as to where they would fall 

in Ruhland’s scheme. In addition, interpretations of several 

species treated by Ruhland have changed. Ruhland (1903) distinguished P. karstenii from P. pilosus by involucral bract pubescence (glabrous vs. pilose), but this is a weak distinction as 

the bracts of P. pilosus are sometimes almost completely glabrous except for the very apex. Similarly, Hensold in Hammel 

et al. (2003a) resurrected P. dendroides (Kunth) Kunth, which 

Ruhland (1903) had considered to be a synonym of P. pilosus. 

The ranges of these two species are similar (Costa Rica south to 

Peru), but there are no records yet of P. dendroides occurring 

in Venezuela. Hensold in Hammel et al. (2003a) distinguished 

P. dendroides from P. pilosus by its larger leaves and peduncles 

and by its glabrous (except for the base) leaves. The leaf margins 

of Guaramacal material of P. pilosus have long cilia (or their 

scars), but leaf length and peduncle length overlap with measurements cited by Hensold in Hammel et al. (2003a) for Costa 

Rican material of P. dendroides. Her brief descriptions in that 

Central American flora do not appear to cover the full range of 

variation in either species.

Heliconiaceae

L. J. DORR AND S. MIGUEL NIÑO

Perennial herbs, terrestrial, solitary or forming dense colonies. Rhizomatous. Aerial stems either very short (leaf sheaths 

forming a pseudostem) or elongate with distinct internodes, enclosed by leaf sheaths. Leaves distichous; composed of sheath, 

blade, and petiole (sometimes inconspicuous or absent); without a distinct ligule; leaf blade entire with a prominent midrib, 

venation generally pinnate-parallel, slightly arcing toward the 

margins. Inflorescences terminal, usually on leafy shoots, sometimes on specialized shoots bearing only bladeless sheaths, with 

few-flowered cymes clustered in axils of cincinnal (or spathaceous) bracts; bracts generally showy (red, yellow, orange or 

a combination of these colors), alternately arranged along the 

rachis, distichous or spiral, persistent or deciduous. Flowers 

bisexual, perfect, 3-merous, heterochlamydeous, strongly zygomorphic. Tepals 6, in 2 series, all petaloid, the median adaxial tepal of outer series distinct, the remaining 5 connate and 

forming a basal tube. Stamens 5, fertile; staminodes 1; filaments adnate to the tubular part of perianth; anthers basifixed, 

2-thecate, 4-sporangiate. Ovary inferior, 3-locular; placentation axile; ovules anatropous, subbasal, 1 per locule; style 1, 

filiform; stigma capitate or 3-lobed. Fruit a drupe; seeds (1–)3, 

not arillate.

A predominately neotropical family composed of a single 

genus and 100–200 species found in Mexico, Central America, 

the West Indies, and South America (all country-level political 

units except Chile and Uruguay); also on the Pacific Islands, including New Guinea. Twenty-three species occur in Venezuela.

Reference. Andersson (1998a).

The majority of Heliconia species found in the Venezuelan 

Andes are known by the common name “platanillo,” which 

reflects the similarity of their leaves to those of the cultivated 

banana or plantain. Some also are known as “ave de paraíso,” 

although less commonly, and this presumably reflects a transfer 

of the common name of the superficially similar Strelitzia Aiton 

(Strelitziaceae), native to Africa and now cultivated throughout 

the tropics.

Many species of Heliconia are cultivated.

MAP 93. Paepalanthus pilosus occurrence in Venezuela.
NUMBER 100 • 9 3

KEY TO THE SPECIES OF HELICONIA

1a. Cincinnal bracts spiral; drupes large, 17–22 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. meridensis

1b. Cincinnal bracts distichous; drupes small, 4–10 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Petioles <1 cm long; flowers resupinate, yellow or orange, always with green eyespots near apex . . . . . . H. hirsuta

2b. Petioles 30–80 cm long; flowers not resupinate, white with a green apex that lacks eyespots . . . . . . . . . . H. stricta

Heliconia hirsuta L. f., Suppl. Pl. 158. 1781 [1782]; Andersson, 

in Berry et al., Fl. Venez. Guayana 5: 585, fig. 504. 1999.

FIGURE 14A–F

Heliconia costanensis Aristeg., Bol. Soc. Venez. Ci. Nat. 25(107): 206, t. 

1964.

Herbs with Zingiber-like habit (i.e., leaves horizontally arranged with short petioles), (0.5–)1.5–3(–4) m tall. Leaves 9–25 

per stem; leaf blades 25–40 × 10–15 cm, glabrous above and 

below, sometimes waxy below; petioles 0.5–1 cm long. Inflorescences erect, to 15(–20) cm long; cincinnal bracts distichous, 

5–10 per inflorescence, basal ones 5–15(–20) × 0.5–1 cm, medial 

ones 4–10 × 0.5–1 cm, usually orange at base, becoming red at 

apex, sometimes greenish-yellow or yellow. Flowers resupinate, 

2.5–5 cm long, yellow to orange or red, with dark green eyespots 

near apex. Ovary 3–6 × 2–4 mm, glabrous to ± hirsute, yellow 

or green with an orange or red apex. Drupes 7–10 × 6–10 mm, 

dark blue.

Found in Central America and South America (Colombia, 

Venezuela, Trinidad and Tobago, the Guianas, Ecuador, Peru, 

Bolivia, Brazil, Paraguay, and Argentina); introduced in the West 

Indies. Widely distributed in Venezuela (Amazonas, Anzoátegui, Apure, Aragua, Barinas, Bolívar, Carabobo, Cojedes, Delta 

Amacuro, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, 

and Zulia; Map 94). In the park, in open areas and in forest 

where the canopy is not completely closed; (1,500–)1,600–1,800

(–2,700) m.

This is a highly polymorphic species.

Heliconia meridensis Klotzsch, Linnaea 20: 462. 1847.

FIGURES 14G–K, 25E

Heliconia schneeana Steyerm., Fieldiana, Bot. 28: 161. 1951.

Heliconia falcata Barreiros, Rodriguésia 28(41): 130. 1976.

Herbs with Musa-like habit (i.e., leaves vertically arranged with long petioles) to Canna-like habit (i.e., leaves 

obliquely arranged with medium-sized petioles), 2–4.5 m tall. 

Leaves 4–5 per stem; leaf blades 100–150 × 25–40 cm, glabrous 

above, ± arachnoid-pilose below on midrib (indumentum often 

Heliconia L.

Heliconia L., Mant. Pl. 2: 147, 211. 1771, nom. cons.

The characters of the genus are those of the family.

References. Andersson (1981b, 1985); Aristeguieta (1961); Berry and Kress (1991); Kress (1997); Kress et al. 

(1999); Rodríguez (1954).

MAP 94. Heliconia hirsuta occurrence in Venezuela. MAP 95. Heliconia meridensis occurrence in Venezuela.
9 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 14. Heliconiaceae. A–F. Heliconia hirsuta L. f. A. Habit. B. Leaf apex. C. Sheath (detail). D. Flower. E. Lateral view of the ovary. 

F. Apical view of the ovary. G–K. H. meridensis Klotzsch. G. Leaf apex. H. Base of leaf (detail). I. Infructescence. J. Lateral view of fruit. K. Apical view of fruit. (A, D, van der Werff et al. 7649; B, C, Cuello et al. 1747; E, F, Dorr et al. 5685; G, Dorr et al. 8341; H, Stergios 17372; I–K, 

R. M. King et al. 10561.)
NUMBER 100 • 9 5

inconspicuous); petioles 15–35(–45) cm long. Inflorescences 

erect, to 33 cm long; cincinnal bracts spiral, 7–9 per inflorescence, 13–15 × 4 cm, generally red, but occasionally displaying 

gradations of pink to dark red. Flowers resupinate, 3–5 cm long, 

yellow or yellowish-green. Ovary 5–6 × 9–10 mm, glabrous, yellow to bright green. Drupes 14–15 × 17–22 mm, dark blue.

Found in South America (Colombia and Venezuela). In Venezuela, found in the Andes (Barinas, Lara, Mérida, Portuguesa, 

Táchira, and Trujillo), the Sierra de Perijá (Zulia), and the Cordillera de la Costa (Yaracuy; Map 95). Common throughout the 

park; (1,500–)1,600–2,000 m.

Heliconia meridensis is sometimes placed in synonymy 

under H. latispatha Benth., but the two species are only superficially similar. The flowers of the former are resupinate (vs. not 

resupinate in the latter), and the perianth is not hidden by cincinnal bracts (vs. partially hidden by cincinnal bracts in the latter).

Heliconia stricta Huber, Bol. Mus. Paraense Emilio Goeldi 4: 

543. 1906. 

Herbs with Musa-like habit, 1.5–4 m tall. Leaves 3–5 per 

stem; leaf blades 80–160 × 15–30 cm, glabrous above and puberulous to villose along midrib margin below; petioles 30–80 cm 

long. Inflorescences erect, 20–35 cm long; cincinnal bracts distichous, 4–8(–10) per inflorescence, basal ones 12–18 cm long, 

apical ones shorter, 8–12 cm long, red or orange on the sides, 

yellowish on keel and margin, green on the edge, rarely entirely 

dingy yellowish. Flowers not resupinate, 5–9 cm long, white basally, dark green distally with white apex. Ovary 5–8 × 3–5 mm, 

glabrous, white. Drupes 11–12 × 4–5 mm, blue.

Found in South America (Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, and Brazil); cultivated in the 

Greater Antilles. In Venezuela, found in Apure, Barinas, Bolívar, 

Mérida, Portuguesa, Táchira, and Trujillo (Map 96). In the park, 

found on the edge of coffee plantations in the karst area above 

the Río Amarillo; ~1,200 m.

Heliconia stricta occurs only near the lower elevational 

limits of the park. It can be confused easily with H. bihai (L.) 

L., which also occurs in the Venezuelan Andes. The former species is less robust and smaller than H. bihai. Also, H. stricta has 

a perianth that is white basally and dark green distally, with a 

white apex (vs. white basally with distal third pale green), and 

the staminodes are inserted (8–)10 mm or usually much more 

above the base of the perianth (vs. 1.5–7 mm above the base of 

the perianth).

Heliconia stricta is widely cultivated and can be seen in 

tropical and subtropical gardens worldwide.

Hypoxidaceae

L. J. DORR

Perennial herbs from ± tuberous rhizomes or corms, the apical portion usually covered by a membranous to fibrous coat 

formed by overlapping leaf bases. Leaves simple, entire, rosulate, 

usually tristichous; blades linear to lanceolate, often plicate (best 

seen in fresh material), pubescent with simple and compound 

hairs, or glabrous, prominently parallel-veined, sessile (our species) or with a petiole-like constriction, persistent. Inflorescences 

several per plant, axillary, borne on leafless scapes arising from 

the tuberous rhizomes or corms, appearing corymbose or racemose, or sometimes reduced to a single flower. Flowers from 

the axils of small or larger and leafy bracts, perfect (our species), 

rarely staminate; tepals usually 6, in 2 whorls, ± petaloid, subequal, free (our species) or fused to form a tube, usually yellow. 

Stamens usually 6, in 1 or less commonly 2 whorls; filaments 

distinct, adnate to base of tepals; anthers 2-loculed, basifixed. 

Ovary inferior, 3-locular; ovules few to many in each locule; 

placentation axile (our species) or parietal; style simple with 3 

stigmas as free lobes or grooves. Nectaries absent. Fruit a capsule with circumscissile or loculicidal dehiscence or sometimes 

indehiscent and berrylike. Seeds small, globose to ellipsoid; testa 

black or brown, smooth to variously textured; embryo small; 

endosperm copious.

A family of 8 genera and 150–175 species widely distributed 

in tropical and subtropical regions of the world, especially in 

the Southern Hemisphere, but also occurring in the Northern 

Hemisphere, most notably in the Americas. In Venezuela, 2 genera (Curculigo Gaertn. and Hypoxis L.) and 3 species are native 

or naturalized. A fourth species, C. capitulata (Lour.) Kuntze, is 

occasionally seen in cultivation, but it is not known to escape in 

Venezuela.

References. Judd (2000); Kocyan et al. (2011); 

Nordal (1998); Rudall et al. (1998).

Morphological and molecular data consistently support Hypoxidaceae as a monophyletic clade (Rudall et al., 1998; Judd, 

MAP 96. Heliconia stricta occurrence in Venezuela.
9 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

2000; Kocyan et al., 2011), which can be subdivided into three 

well-supported major clades (Kocyan et al., 2011). Genera and 

species belonging to two of these three major clades, the Curculigo clade and the Hypoxis clade, are found in the Venezuelan 

flora.

Hypoxis L.

Hypoxis L., Syst. Nat., ed. 10, 972, 986, 1366. 1759.

Rhodohypoxis Nel, Bot. Jahrb. Syst. 51: 239, 257, 300. 1914.

Acaulescent herbs; rhizomes often relatively large and ± 

tuberous. Leaves grasslike, linear-lanceolate to almost filiform, 

usually pubescent, even if only sparingly so. Inflorescences reduced cymes of 2 to several flowers, but often appearing racemose, sometimes reduced to a solitary, axillary flower, borne on 

a slender, erect to decumbent scape that is usually shorter than 

the subtending leaves. Flowers sessile to long-pedicellate and of 

2 kinds: chasmogamous flowers showy, with spreading tepals, 

cleistogamous ones inconspicuous, with perianth parts forming 

a conical point. Tepals ± petaloid, 3 outer tepals spreading, narrower than inner, green below and yellow above, inner tepals 

spreading, yellow above and below (white to green in cleistogamous flowers). Stamens exserted, the filaments short subulate to 

filiform. Fruit a capsule dehiscing by a circumscissile slit below 

the marcescent perianth and often then splitting longitudinally 

into 3 segments; seeds spherical-ellipsoid.

A genus of 50–100 species widely distributed in America, 

Africa, Madagascar, Asia, and Australia, with an important center of diversity in Africa. Species number is problematic since the 

genus is apomictic. Two species occur in Venezuela.

Reference. Brackett (1923).

Problems with circumscribing a monophyletic Hypoxis

were noted by Judd (2000). Molecular data (Kocyan et al., 2011) 

suggest that such a circumscription can be achieved by placing 

the African genus Rhodohypoxis Nel in synonymy (as is done 

here) and by excluding several Australian species of Hypoxis

that show affinities to the one major clade of Hypoxidaceae not 

found in Venezuela and otherwise restricted to southern Africa.

Mature seeds are needed to identify specimens as the 

color and ornamentation of the testa are important diagnostic 

characters.

Hypoxis decumbens L., Syst. Nat., ed. 10, 986. 1759; Morillo, 

in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 

510–511, foto 1. 2010.

FIGURE 15

Hypoxis decumbens var. major Seub., in Martius, Fl. Bras. 3(1): 52, t. 7, fig. 

1. 1847.

Herbs 10–30(–45) cm tall. Leaves linear to linear-lanceolate, 

5–30(–45) × 0.2–0.8 cm, sparingly pubescent with long, simple 

hairs, especially along the margin; leaf sheaths disintegrating 

completely, not fibrous. Inflorescences 3–5 per plant, each scape 

with 1–4(–6) flowers, scapes filiform, (5–)10–20(–30) cm long, 

shorter than the leaves, sparingly pubescent to glabrous. Tepals 

ovate-lanceolate, 4–6 × 1.5–3 mm, glabrous adaxially, pubescent 

abaxially. Capsules subcylindric to fusiform, 10–15(–20) × 2–3 

mm, densely pubescent to glabrate. Mature seed black, ellipsoid, 

~1 mm in diameter; testa tuberculate.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Ecuador, including the 

Galapagos Islands, Peru, Bolivia, Brazil, Paraguay, Argentina, 

and Uruguay). In Venezuela, found in the Andes (Lara, Mérida, 

Miranda, Portuguesa, Táchira, and Trujillo) and in the Cordillera de la Costa (Aragua, Distrito Federal, and Yaracuy; Map 

97). We have a few collections from below Páramo del Pumar 

and in El Campamento on the south slope (Qda. Jirajara), but 

we suspect that this species has a wider distribution in the park 

in disturbed areas and pastures; 1,900–2,800 m.

Use. A glue is obtained from the rhizomes (R. Caracas, pers. comm.).

Hypoxis decumbens can be distinguished from H. humilis

Kunth, which also occurs in Venezuela, by seed color (black vs. 

dull brown) and texture (rounded tubercles vs. sharp-pointed 

murications). We do not agree with Campbell in Hokche et al. 

(2008: 744), who synonymized these two species, and we have 

not incorporated her distributional data into ours. Undoubtedly, 

H. decumbens occurs in more Venezuelan states than we have 

indicated here.

Iridaceae

L. J. DORR AND B. STERGIOS

Perennial or rarely annual herbs, woody shrubs (Africa only) 

or achlorophyllous saprophytes (Madagascar only); sometimes 

MAP 97. Hypoxis decumbens occurrence in Venezuela.
NUMBER 100 • 9 7

caespitose; underground stems rhizomes, corms or bulbs. Leaves 

basal and cauline, sometimes cataphylls present, distichous, 

sheaths of the foliage leaves open or closed; leaf blades unifacial, 

but bifacial proximally (equitant) or occasionally terete, plane or 

plicate, venation parallel. Flowering stems simple or branched, 

terete or compressed and then angled or winged. Inflorescences 1 

to many umbellate monochasial cymes (rhipidia), panicles, spikes 

or flowers solitary; rhipidia enclosed in 2, opposed, large, leafy 

bracts (spathes). Flowers usually pedicellate (or subsessile), bisexual, actinomorphic to slightly zygomorphic; perianth petaloid 

FIGURE 15. Hypoxidaceae, Hypoxis decumbens L. A. Habit. B. Leaf (detail of abaxial surface). C. Flower. D. Flower, pistil removed. E. Mature fruit. F. Seed. (A–F, Dorr et al. 8556.)
9 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

with 2 equal or unequal whorls of 3 tepals each; tepals showy, 

distinct or connate below to form a tube. Stamens 3(rarely 2), 

opposite the outer tepals; filaments distinct or connate below 

to form a tube; anthers 2-thecate, 4-sporangiate, extrorse or 

latrorse, usually dehiscing by longitudinal slits. Ovary inferior 

(rarely superior), 3-locular; placentation axile; ovules many to 

few (occasionally 1 or 2), anatropous; style single below, filiform 

(at least proximally), 3-branched or 3-lobed, branches either filiform or distally expanded or branches thickened or flattened, 

petaloid. Fruits capsular, loculicidal, rarely indehiscent. Seeds 

globose to angular or discoid; seed coat reticulate to smooth, 

generally dry, rarely fleshy or arillate.

A cosmopolitan family of 65–75 genera and 2,000–2,050 

species, with southern Africa the most important center of diversity. The family is found throughout the Americas. In Venezuela, 

12 genera and ~20 species of native or naturalized taxa have 

been recorded.

References. Goldblatt and Manning (2008); Goldblatt et al. (1998, 2008); Reeves et al. (2001a, 2001b).

Iridaceae are popular garden plants, and additional genera 

and species are likely to become naturalized in Venezuela.

KEY TO THE GENERA OF IRIDACEAE

1a. Flowering stalks terete; flowers ± sessile, included within the spathes; capsules pubescent; seeds angular, whitish-brown 

to brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orthrosanthus

1b. Flowering stalks compressed, usually angular or winged; flowers long-pedicellate, exserted from the spathes; capsules 

glabrous; seeds globose or compressed-globose, dark brown to blackish . . . . . . . . . . . . . . . . . . . . . . . . . . Sisyrinchium

Orthrosanthus Sweet

Orthrosanthus Sweet, Fl. Australas. t. 11. 1827.

Perennial, evergreen herbs; caespitose and forming large 

tufts; rhizomes short, persistent, creeping. Leaves mostly basal, 

narrowly ensiform, equitant, linear to linear-lanceolate, coriaceous, without a prominent midrib. Flowering stems erect, 

branched or semibranched, branches long or short, ascending, terminating in a single rhipidium or branches with various rhipidia; rhipidia ± short; spathes (bracts) unequal. Flowers 

bisexual, actinomorphic, subsessile or short-pedicellate, fugacious. Tepals free (our species) or connate in a short tube (Australian species), subequal, spreading horizontally from the base, 

blue or white. Anthers erect; filaments free or partially connate 

below. Ovary subsessile, included within the spathes (our species), rarely exserted; style short, 3-branched, branches relatively 

long, extending between the stamens, and stigmatic apically. 

Capsule ellipsoid to cylindric, pubescent (our species) or not. 

Seeds angular to fusiform.

A genus of 9 species with an unusual disjunct distribution; 

5 species found in Mexico, Central America, and South America 

(Colombia, Venezuela, Ecuador, Peru, Bolivia, and Argentina) 

and 4 in southwestern Australia. The American species are restricted to higher elevations but are absent from high elevations 

in the Venezuelan Guayana. Two species occur in Venezuela.

References. Henrich and Goldblatt (1987); Steyermark (1948).

Orthrosanthus is superficially similar to Eccremis Willd. ex 

Baker (Xanthorrhoeaceae). Characters to distinguish the two 

genera in Guaramacal are listed following the description of 

Eccremis.

Orthrosanthus acorifolius (Kunth) Ravenna, Not. Mens. Mus. 

Nac. Hist. Nat. 21(249): 9. 1977; Morillo, in Morillo et 

al., Bot. Ecol. Monocot. Páramos Venez. 2: 515–516, fig. 1. 

2010. Moraea acorifolia Kunth, in H. B. K., Nov. Gen. Sp. 

[fol.] 1: 259. 1815 [1816]; ibid. [qu.] 1: 322. 1815 [1816]. 

Orthrosanthus chimboracensis var. acorifolius (Kunth) 

Steyerm., Lloydia 11: 19. 1948.

FIGURE 16D–G

Orthrosanthus chimboracensis auct., non (Kunth) Baker; Ortega et al., BioLlania 5: 43. 1987.

Robust herbs, 0.8–1 m tall. Basal leaves 40–50 × 1.5–2 cm, 

broadly linear, gradually acuminate, sometimes plicate, cauline 

leaves 10–20 cm long, linear-acuminate. Flowering stems erect, 

rigid, branched with internodes, ± equal in length to basal leaves, 

0.5 cm wide. Rhipidia several-flowered, flowers ~3 cm in diameter, 

MAP 98. Orthrosanthus acorifolius occurrence in Venezuela.
NUMBER 100 • 9 9

showy, blue or bluish-white. Capsules 8–11 mm long, sessile or 

short-pedicellate, pubescent. Seeds rounded-angular or fusiform.

Found in South America (Colombia and Venezuela). In Venezuela, found in the Andes (Lara, Mérida, and Trujillo) and in the 

Cordillera de la Costa (Anzoátegui, Distrito Federal, Miranda, 

and Sucre; Map 98). Collected in páramo and subpáramo in the 

park, including the Fila de Agua Fría, the Páramo de Guaramacal, and the Páramo de Vicuyal and between El Campamento 

and El Santuario; (1,900–)2,100–3,100 m.

Common Names. Espadilla and palmiche.

FIGURE 16. Iridaceae. A–C. Sisyrinchium tinctorium Kunth. A. Habit. B. Flower showing ovary. C. Capsule. D–G. Orthrosanthus acorifolius

(Kunth) Ravenna. D. Habit. E. Open flower. F. Flower showing ovary. G. Pubescent capsule. (A, Aymard et al. 4409; B, C, Aymard et al. 5180; 

D, Stergios 17469; E, F, Stergios 18723; G, Cuello 1387.)
100 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Orthrosanthus acorifolius is distinguished from O. chimboracensis (Kunth) Baker, the only other species found in the Venezuelan Andes, by its sessile or shortly pedicellate (pedicels <5 

mm long) and pubescent fruit, by its smaller stature, and by the 

fact that its flowering stalks scarcely exceed the leaves in length.

The label of one collection from Guaramacal (Cuello 1387) 

states that the flowers are yellow, which is without doubt an error.

Sisyrinchium L.

Sisyrinchium L., Sp. Pl. 954. 1753.

Perennial or rarely annual herbs; often caespitose; rhizomatous or rootstock indistinct, roots thick and fleshy or tuberous or 

fibrous (annual species). Leaves basal or basal and cauline, basally equitant; leaf blades ensiform, lanceolate to linear, flat (our 

species) or sometimes terete, usually glabrous. Flowering stems 

scape-like or branched, compressed and angled or winged, sometimes comprising a single extended terminal internode; spathes 

equal or subequal. Flowers actinomorphic; tepals ± distinct, ± 

equal, rotate or campanulate, yellow, white or blue with a yellow 

center, sometimes with darker-colored veins. Stamens symmetrically arranged; filaments distinct, connate basally (our species) 

or united in a tube, often puberulent or glandular-pubescent. 

Ovary globose, included or exserted; style 3-branched, suberect, 

connate below, filiform, branches extending between stamens. 

Capsule globose to cylindric or pyriform, glabrous (our species) 

or papillose. Seeds globose and slightly flattened or 3-angled, frequently foveolate or deeply pitted.

An almost exclusively American genus of 100–150 species 

with a wide distribution, including North America (Greenland, 

Canada, and USA), Mexico, Central America, the West Indies, 

and South America (Colombia, Venezuela, Guyana, Ecuador, Peru, 

Bolivia, Brazil, Chile, Paraguay, Argentina, and Uruguay); also, 1 

species is native to Hawaii. Species of Sisyrinchium are naturalized 

in the Galapagos Islands, Europe (western Ireland), the Mascarene 

Islands, Malaysia, and a number of the Pacific Islands, including 

Hawaii and New Zealand. Four species occur in Venezuela.

References. Chauveau et al. (2011); Goldblatt et 

al. (1990); Karst and Wilson (2012); Tacuatiá et al. (2012).

Sisyrinchium is the largest genus of Iridaceae in the Americas.

KEY TO THE SPECIES OF SISYRINCHIUM

1a. Flowering stem branched, or at least with 1 or more cauline leaves and several internodes, not or minutely winged, 

1 mm wide; flowers white or lilac-pink, with dark maroon veins; rhipidium sometimes repeated in second-order branches

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. micranthum

1b. Flowering stem unbranched, a single long, leafless internode, narrowly to broadly winged, 3 mm wide; flowers pale 

yellow, without dark maroon veins; rhipidium solitary and terminal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tinctorium

Sisyrinchium micranthum Cav., Diss. 6: 345, t. 191, fig. 2. 

1788; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 517–518, fig. 2. 2010.

Sisyrinchium iridifolium Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 260. 1815 

[1816]; ibid. [qu.] 1: 324. 1815 [1816].

Delicate, annual herbs, 6–25 cm tall; roots fibrous. Leaves 

linear, acuminate, basal leaves 5–9 cm × 2–4 mm, cauline 

leaves smaller, 1.5–6 cm × 1–2 mm. Flowering stems branched, 

4–20 cm tall, 0.8–1 mm wide, not or minutely winged, with several internodes with cauline leaves; spathes 1.5–2(–4) cm long. 

Flowers campanulate, white or lilac-pink and with dark maroon 

veins (our material), blue with a yellow center or yellow; tepals 

6–7.5 mm long. Capsules globose, ~3 mm long. Seeds minute, 

<1 mm in diameter, 3-angled, pitted, blackish.

Native to Mexico, Central America, the Greater Antilles 

(Hispaniola), and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Paraguay, and Argentina); adventive 

in the Galapagos Islands, the Mascarene Islands, and several Pacific Islands, including New Zealand. In Venezuela, found in the 

Andes (Mérida, Táchira, and Trujillo) and in the Cordillera de la 

Costa (Aragua, Distrito Federal, and Miranda; Map 99). In the 

park, found in open and rocky areas at the edge of forest on both 

slopes of Guaramacal; (1,700–)2,000–2,500 m.

This is a morphologically and cytologically complex species 

(Tacuatiá et al., 2012) with an extensive range and a tendency 

to be invasive.

Sisyrinchium tinctorium Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 

1: 260. 1815 [1816]; ibid. [qu.] 1: 324. 1815 [1816]; Goldblatt and Henrich, in Berry et al., Fl. Venez. Guayana 5: 

MAP 99. Sisyrinchium micranthum occurrence in Venezuela.
NUMBER 100 • 101

662. 1999; Morillo, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 2: 518–519. 2010.

FIGURES 16A–C, 25G

Sisyrinchium bogotense Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 259. 1815 

[1816]; ibid. [qu.] 1: 323. 1815 [1816].

Perennial herbs, (15–)22–32 cm tall; roots mostly fibrous, 

some thickened and tuberous, but always thin. Leaves all basal, 

linear-lanceolate, 18–25 cm × 5–7(–12) mm, acuminate. Flowering stems unbranched, equal to or scarcely surpassing the 

leaves in length, 12–24 cm tall, 3 mm wide, narrowly to broadly 

winged; spathes 3–4 cm long. Flowers stellate, pale yellow, without dark maroon veins; tepals 7–10 mm long. Capsules pyriform, 10–15 mm long, pendulous. Seeds ~1.5 mm in diameter, 

globose, foveolate, blackish.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Ecuador, Peru, Bolivia, and Argentina). 

In Venezuela, found in the Andes (Lara, Mérida, Táchira, and 

Trujillo), the Cordillera de la Costa (Aragua, Miranda, Monagas, and Sucre), and possibly Amazonas (Map 100). Open areas 

in montane forest, cloud forest, and especially páramo in the 

park; 1,800–2,800 m.

Common Name. Espadilla.

The record from Amazonas state (Goldblatt and Henrich in 

Berry et al., 1999: 662) is anomalous as it is outside the expected 

range of this species. The specimen upon which the record is 

based has not been examined by us.

Juncaceae

L. J. DORR AND S. MIGUEL NIÑO

Perennial or rarely annual herbs; terrestrial or sometimes 

rooted aquatics; usually rhizomatous or caespitose. Flowering 

stems erect, ascending or rarely procumbent, terete or laterally 

compressed; leafless or leaf-bearing. Leaves simple, spirally arranged, tristichous or rarely distichous, basal or on lower part of 

stem, blades linear to filiform, flat or terete, when terete hollow 

and sometimes septate, margins glabrous or pilose (Luzula DC.); 

leaf sheaths open or closed (Luzula), auriculate or not; blades of 

basal leaves sometimes reduced (cataphylls). Inflorescences terminal or pseudolateral, compound, cymose or racemose, paniclelike or anthelate (i.e., a panicle in which the lower branches 

overtop the upper ones), many-flowered, flowers in 1 to many 

terminal heads or spikelike clusters, or flowers sometimes solitary 

and terminal or lateral; lower inflorescence bracts usually herbaceous; flower bracteoles 1 or 2 or absent. Flowers perfect, rarely 

dioecious or monoecious, actinomorphic, hypogynous. Tepals 6 

in 2 whorls, free, equal or subequal, glumaceous or herbaceous. 

Stamens (2–)3–6, in 2 whorls, alternate to the tepals; anthers basifixed, 2-locular, oblong or linear. Ovary superior, 1-locular or 

3-septate to 3-locular; ovules 3 to many, central, basal or ± parietal, anatropous; style 1-, 2-, or 3-branched (stigmas). Fruit a 

dry capsule, loculicidal or rarely circumscissile. Seeds 3 to many, 

globose, angled or flattened, sometimes with tail-like appendages.

A cosmopolitan family of 7 genera and 400–450 species 

generally found in cool and temperate regions. The family is 

found throughout the Americas but is absent from the Guianas. 

Two genera and 14 species occur in Venezuela.

References. Balslev (1996, 1998); Drábková et al. 

(2003); Kirschner (2002a, 2002b, 2002c); Záveská Drábková 

(2010); Záveská Drábková and Vlček (2009).

The circumscription of most of the genera of this family is 

likely to change as additional molecular evidence is gathered. Although Luzula appears to be monophyletic, Juncus L., the largest genus, is clearly paraphyletic (Drábková and Vlček, 2009; 

Záveská Drábková, 2010); the small South American genera Distichia Nees & Meyen, Marsippospermum Desv., Oxychloe Phil., 

Patosia Buchenau, and Rostkovia Desv. (Marsippospermum and 

Rostkovia are also found in New Zealand and subantarctic islands) are embedded in a clade that includes species of Juncus.

Prionium E. Mey., a small genus endemic to southern Africa, 

often has been included in the Juncaceae, but molecular data 

indicate that it is sister to Juncaceae and Cyperaceae (Munro 

and Linder, 1998; Davis et al., 2004) and is better placed in the 

closely related Thurniaceae (synonym Prioniaceae), a family of 

two genera with an unusual southern African–South American 

disjunction.

KEY TO THE GENERA OF JUNCACEAE

1a. Leaf margins glabrous; leaf sheaths open (rarely closed); capsules 1- or 3-locular; seeds many . . . . . . . . . . . . . . . Juncus

1b. Leaf margins (at least when young) pilose; leaf sheaths closed; capsules 1-locular; seeds 3 . . . . . . . . . . . . . . . . . Luzula

MAP 100. Sisyrinchium tinctorium occurrence in Venezuela.
102 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Juncus L.

Juncus L., Sp. Pl. 325. 1753.

Perennial or rarely annual herbs; rhizomatous or caespitose, 

glabrous. Stems usually erect, sometimes prostrate or ascending. Basal leaves lacking blades or blades reduced (cataphylls); 

fully developed leaves basal and/or cauline, spiraled, flat and not 

septate or terete and with or without septa; leaf sheaths open, 

usually conspicuously auriculate or without auricles. Bracts leaflike or reduced, decreasing in size from base to apex. Inflorescences compound, cymose or racemose, often anthelate, with 

few to many flowers, sometimes clustered in heads. Flowers bisexual (rarely unisexual), with or without a pair of bracteoles. 

Tepals lanceolate, equal or nearly equal, entire, persistent. Stamens (2–)3–6; filaments filiform or flattened; anthers linear to 

oblong. Ovary sessile; style almost absent to long; stigmas (2) 

3, filiform, twining, sticky papillose. Capsules 1- or more commonly 3-locular. Seeds numerous, ellipsoid to ovoid, appendaged 

or not.

As presently construed a cosmopolitan genus of ~300 species, ~50 of which are tropical and confined to higher (>1,500 

m) elevations. Juncus occurs in all major political units in the 

Americas with the exception of the Guianas. Twelve species 

occur in Venezuela.

References. Kirschner (2002b, 2002c).

Molecular evidence suggests that Juncus is paraphyletic 

(Záveská Drábková and Vlček, 2009; Záveská Drábková, 2010); 

the majority of taxa demonstrate affinity to two main groups on 

the basis of the presence or absence of bracteoles and cymose or 

racemose inflorescences, respectively. However, many species lie 

outside this scheme, and there are also species where bracteoles 

may be present or not.

KEY TO THE SPECIES OF JUNCUS

1a. Annuals; stems scapose; leaf blades flat, never septate; leaf sheaths without auricles; inflorescences open or pseudolateral; 

flowers mainly solitary, each flower with 2 clasping bracteoles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. bufonius

1b. Perennials; stems ± leafy; leaf blades cylindric, septate; leaf sheaths with auricles; inflorescences capitate; flowers clustered, 

without clasping bracteoles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. microcephalus

Juncus bufonius L., Sp. Pl. 328. 1753; Briceño, in Morillo et 

al., Bot. Ecol. Monocot. Páramos Venez. 2: 523–524, fig. 

2. 2010.

Annual herbs, 5–35 cm tall; caespitose; without rhizomes. 

Stems erect, procumbent or ascending, terete, smooth. Basal 

leaves often cataphylls; cauline leaves linear, plane or involute 

(edges incurved toward the top of the leaf) with raised margins, 

slightly channeled, to 20 cm × 0.5–1 mm; sheaths 1–3 cm long, 

auricles absent. Inflorescences compound, lax, cymose; lowermost inflorescence bracts always the longest, 10–15 cm long; 

uppermost bracts to 5 mm long; cymes with solitary flowers, 

sometimes with 2–4 flowers, each flower subtended by a pair of 

bracteoles. Tepals unequal, lanceolate, acuminate, green. Capsules ellipsoid, truncate, 3.5–5 × 2 mm.

A cosmopolitan weed found only in high elevations in the 

tropics, typically in open and wet sites. Its American distribution 

is extensive and includes North America, Mexico, Central America, the West Indies, and South America (Colombia, Venezuela, 

Ecuador, Peru, Bolivia, Brazil, Chile, Argentina, and Uruguay). 

In Venezuela, known from the Andes (Mérida, Táchira, and Trujillo) and the Cordillera de la Costa (Aragua, Distrito Federal, 

and Monagas; Map 101). In the park, known from the Páramo 

de Guaramacal and elsewhere along the Boconó–Guaramacal 

road; 1,600–3,100 m.

This species is part of “an intricate complex of very similar 

and closely related taxa” (Kirschner, 2002c: 9).

Juncus microcephalus Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 

190. 1815 [1816]; ibid. [qu.] 1: 237. 1815 [1816]; Briceño, 

in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 

527, fig. 6. 2010.

FIGURE 17C–H

Juncus densiflorus auct., non Kunth; Dorr et al., Contr. U.S. Natl. Herb. 40: 

47. 2000 [2001].

Perennial herbs, 20–75(–100) cm tall; caespitose; rhizomes 

short, densely branched. Stems erect, terete, smooth or striate. 

Leaf blades linear, terete, hollow, striate and transversely septate, 

MAP 101. Juncus bufonius occurrence in Venezuela.
NUMBER 100 • 103

FIGURE 17. Juncaceae. A, B. Luzula gigantea Desv. A, A1. Habit. B. Inflorescence (detail). C–H. Juncus microcephalus Kunth. C. Habit.

D. Head. E. Flower with ovary removed showing stamens and tepals. F. Flower with three tepals and ovary removed showing three stamens. 

G. Flower with tepals removed showing stamens, ovary, and styles. H. Cross section through leaf blade. (A, B, Stergios et al. 20549; C, Stergios 

& Zambrano 17686; D–H, Cuello et al. 1266.)
104 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

10–40 cm × 2–3 mm; sheaths 4–12 cm long, auriculate. Inflorescences anthelate, decompound, composed of conical to globose heads, heads 2–6-flowered; the lowermost bracts (of the 

peduncle) usually sheath-like, 5–10 cm long, generally shorter 

than the inflorescence. Tepals unequal, lanceolate, acute, cleargreen to pale coffee. Capsules ellipsoid to obovoid, truncate, 2–3 

× 1.5 mm.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Chile, Paraguay, Argentina, and Uruguay); introduced in Australia. In Venezuela, found in the Andes (Lara, Mérida, Portuguesa, Táchira, 

and Trujillo), the Sierra de Perijá (Zulia), and the Cordillera de la 

Costa (Distrito Federal and Sucre; Map 102). In the park, found 

on both slopes of Guaramacal, including near the summit of the 

Fila de Agua Fría; (1,500–)1,600–3,100 m.

Common Name. Paja grama.

This species evidently is absent from the Venezuelan Guayana despite a record for Amazonas state (Schnee in Pittier et al., 

1945: 172), which should be treated with skepticism.

Luzula DC.

Luzula DC., in Lamarck and A. P. de Candolle, Fl. Franç., 3rd ed., 3: 158. 

1805, nom. cons.

Perennial or rarely annual herbs; usually caespitose; rhizomatous. Stems erect. Leaves alternate, scalelike on stolons and rhizomes, leafy and grasslike at the base of the stem, and bract-like 

below the inflorescence. Leaf blades linear to narrowly lanceolate, plane, margins pilose (at least when young); sheaths closed, 

auricles absent. Inflorescences diverse, usually many-flowered, 

either cymose with flowers borne singly or in few-flowered 

clusters in much-branched panicles or decompound anthelate 

inflorescences, or in spikelike clusters, either pedunculate or 

clustered in many-flowered dense heads. Flowers perfect, each 

flower subtended by a small bract at pedicel base, and 1 or 2 

bracteoles just below flower; bracts decreasing in size from the 

base to the apex and changing in color from green to chestnut or 

bright brown. Tepals 6, equal to ± unequal, lanceolate, margin 

entire (our species) or ciliate. Stamens 3–6; anthers oblong to 

linear; filaments filiform, rarely linear or flattened. Ovary sessile; 

style short, filiform; stigmas 3, filiform, twining, sticky papillose. 

Capsules 1-locular, ± ovoid-trigonous, occasionally 3-lobed basally. Seeds 3, with or without appendages.

A cosmopolitan genus of ~115 species found principally in 

temperate regions. Approximately 12 species occur in tropical 

regions of the world, generally at higher (>2,500 m) elevations. 

The genus is present throughout the Americas, with the exception of parts of Central America, the West Indies, and the Guianas, Paraguay, and Uruguay in South America. Only 2 species 

have been collected in Venezuela, both in the Andes.

Reference. Kirschner (2002a).

Luzula as traditionally circumscribed appears to be monophyletic (Drábková and Vlček, 2009; Záveská Drábková, 2010).

Luzula gigantea Desv., J. Bot. (Desvaux) 1: 145. 1808; Briceño, 

in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 

529–530, fig. 8. 2010.

FIGURE 17A,B

Perennial herbs, 15–90 cm tall; rhizomatous. Stems erect, 

terete, 2–4 mm in diameter, ± rosulate. Basal leaves 10–30 ×1–2 

cm, linear to narrowly lanceolate, plane, margins pilose (at least 

when young); sheaths short; cauline leaves similar to basal ones, 

MAP 103. Luzula gigantea occurrence in Venezuela.

MAP 102. Juncus microcephalus occurrence in Venezuela.
NUMBER 100 • 105

but sheaths to 6 cm long. Inflorescences lax, compound panicles, 

storied and subdivided into 4–5 subequal anthelate parts; flowers solitary on elongate and distinct pedicels; bracteoles ovate to 

ovate-lanceolate. Tepals ± unequal. Stamens 6. Capsules widely 

ellipsoid, apiculate, ~2 × 1.5 mm. Seeds ~1 × 0.5 mm, appendages scarcely visible.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, Bolivia, and Argentina). In Venezuela, known only from 

the Andes (Mérida, Táchira, and Trujillo; Map 103). In the park, 

found in páramo near the Laguna del Pumar; 2,900–3,100 m.

Luzula racemosa Desv., the only other species of the genus 

found in Venezuela, can be distinguished by its spiciform raceme.

Marantaceae

L. J. DORR AND S. MIGUEL NIÑO

Perennial herbs, terrestrial (our species) or semiaquatic; 

rhizomatous; stems soft to suffruticose, usually branched, sometimes giving the plant a shrubby appearance. Leaves in a basal 

cluster, both basal and cauline or solely cauline, alternate, distichous, consisting of an open sheath, petiole (occasionally absent), and blade; ligule absent; blade with a prominent midrib 

and parallel veins, lateral veins ± sigmoid and united near the 

margin of the blade; petiole modified at apex as a pulvinus. Inflorescences 1 to several per shoot (synflorescence), varying in 

level of complexity; each inflorescence terminal (or terminal 

and lateral), usually a spicate thyrse with bracteate spathes subtending groups of 1 to several cymules of 2 flowers each; the 

spathes distichous or spirally arranged, usually showy and colored; bracteoles present or absent. Flowers bisexual, epigynous, 

asymmetrical (the pair of flowers of each cymule appearing as 

mirror images). Sepals 3, distinct, not petaloid. Petals 3, 1 often 

cucullate and larger than the other 2. Corolla, staminodes, and 

style fused and forming a tube; fertile stamen 1, sometimes the 

filament petaloid; anthers 1-thecate; staminodes (2–)3–4, outer 

staminodes (0) 1 or 2, petaloid, inner staminodes 2, 1 hooded 

(staminodium cucullatum) and 1 firm and fleshy (staminodium 

callosum), appendaged. Ovary inferior, 3-locular or 1-locular 

by abortion; placentation basal; septal nectaries well-developed. 

Ovule 1 per locule, anatropous. Style 1, terminal portion and 

stigma enclosed in the hooded staminode (staminodium cucullatum) and held under tension until explosively released by a 

pollinator; stigma cup-shaped. Fruit a capsule, berry or achene, 

dehiscent or rarely indehiscent, fleshy or dry; sepals often persistent in fruit. Seeds 1–3 with a basal aril or aril absent.

A family of 31 genera and ~550 species distributed in the 

tropics and subtropics of the Americas, Africa, Madagascar, and 

Asia. In Venezuela there are 9 genera and 62 species.

References. Andersson (1976, 1998b); Andersson 

and Chase (2001); Prince and Kress (2006); Rogers (1984).

Andersson (1976) provides a detailed discussion of the complicated inflorescence structure of Marantaceae.

The majority of Venezuelan species of Marantaceae are 

known by the common name “platanillo,” which is derived from 

the similarity of their leaves to those of the cultivated banana or 

plantain.

A number of species of Marantaceae are cultivated as ornamental plants, especially for their foliage. Also, the tubers of at 

least one species found in Venezuela, Maranta arundinacea L., 

are the source of an edible starch.

Stromanthe Sond.

Stromanthe Sond., Neue Allg. Deutsche Garten-Blumenzeitung 5: 225. 1849.

Shrub-like herbs, generally caulescent. Leaves clustered 

at the nodes, petiolate, with pulvinus or not. Inflorescences 

terminal, usually well branched, floral units composed of 1–5 

two-flowered cymules; bracteate spathes green or bright colors 

(orange, yellow or red), deciduous or not; bracteoles present or 

absent. Sepals fibrous. Tube formed by fusion of petals, stamens, 

staminodes, and discrete style shorter than wide. Petals blueviolet (lavender), white or greenish. Ovary with 2 sterile locules 

and 1 fertile one. Capsules with a thick pericarp, sepals persistent or not. Seed 1, arillate.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Trinidad and Tobago, the 

Guianas, Ecuador, Peru, Bolivia, Brazil, and Argentina); ~15 species. In Venezuela, only the 2 species treated here are found.

Reference. Andersson (1981a).

KEY TO THE SPECIES OF STROMANTHE

1a. Leaves 20–40 × 4.5–16 cm, pulvinus 1.5–2.5(–3) cm long; spathes showy, orange or bright yellow; petals white to greenish; ovary glabrous or slightly pubescent, fruit yellowish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. jacquinii

1b. Leaves 5–20 × 2.5–5 cm, pulvinus 0.4–1 cm long; spathes not showy, light green; petals white to lavender; ovary densely 

pubescent; fruit orange to reddish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tonckat

Stromanthe jacquinii (Roem. & Schult.) H. Kenn. & Nicolson, Ann. Missouri Bot. Gard. 62: 501. 1975; Andersson, 

in Berry et al., Fl. Venez. Guayana 6: 247. 2001. Maranta 

jacquinii Roem. & Schult., Syst. Veg. 1: 558. 1817 (“jacquini”).

Maranta lutea Jacq., Ic. Pl. Rar. 2: 1, t. 201. 1788, non Aubl., 1775.

Shrub-like herbs, to ~2 m tall. Leaf blades 20–45 × 4.5–16 

cm, with hairs along the midrib above and below; pulvinus well 

developed, 1.5–2.5(–3) cm long, slightly tomentose basally. Inflorescences monosymmetric; spathes bract-like, showy, orange 

or bright yellow. Petals white to greenish. Ovary glabrous or 

slightly pubescent. Fruit yellowish.
106 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Found in Central America (Panama) and South America 

(Colombia, Venezuela, and Ecuador); adventive in Hawaii. In 

Venezuela, found in the Andes (Barinas, Lara, Mérida, Portuguesa, Táchira, and Trujillo), the Sierra de Perijá (Zulia), the Cordillera de la Costa (Aragua, Carabobo, Distrito Federal, Falcón, 

Miranda, and Yaracuy), and the Venezuelan Guayana (Bolívar; 

Map 104). Collected in forest understory near the lower limits 

of the southern boundary of the park in the karst area above the 

Río Amarillo; ~1,200–1,400 m.

Stromanthe tonckat (Aubl.) Eichler, Abh. Königl. Akad. Wiss. 

Berlin 1883: 80. 1884; Andersson, in Berry et al., Fl. Venez. 

Guayana 6: 247. 2001. Maranta tonckat Aubl., Hist. Pl. 

Guiane 1: 3. 1775.

FIGURE 18

Shrub-like herbs, to 3 m tall. Leaf blades 5–20 × 2.5–5 cm, 

glabrous below and with hairs along the midrib above; pulvinus 

short, 0.4–1 cm long, tomentose basally. Inflorescences bisymmetric, greatly reduced; spathes bract-like, greenish. Petals lavender, gradually changing to white. Ovary densely pubescent. Fruit 

orange to reddish.

Found in Mexico, Central America, the Lesser Antilles, and 

South America (Colombia, Venezuela, Trinidad and Tobago, the 

Guianas, Bolivia, and Brazil). In Venezuela, recorded from Anzoátegui, Apure, Aragua, Barinas, Bolívar, Distrito Federal, Falcón, 

Guárico, Mérida, Miranda, Nueva Esparta, Portuguesa, Sucre, 

Táchira, Trujillo, and Yaracuy (Map 105). In the park, found 

only at lower elevations on the south slope in localities such as 

La Divisoria de la Concepción and the karst area above the Río 

Amarillo; ~1,200–1,500 m.

Orchidaceae

G. CARNEVALI FERNÁNDEZ-CONCHA AND L. J. DORR

Terrestrial, lithophytic or epiphytic, rarely subaquatic or 

subterraneous herbs, or rarely vines; chlorophyllous (our species) 

or rarely achlorophyllous; strongly mycotrophic. Rhizomes delicate to stout, sometimes creeping, simple or branched. Roots fibrous or fleshy to tuberous, in epiphytic taxa frequently sheathed 

by 1 or several layers of velamen (i.e., water-absorbing layer of 

dead cells) originating from the rhizome or from the bases or 

sides of stems. Stems elongate to almost absent, primarily with a 

sympodial branching pattern, less commonly monopodial; stems 

in sympodial systems frequently thickened into corms or pseudobulbs. Leaves with parallel venation, simple (rarely lobed), 

articulate or not with their sheaths, distichous or spiraled, membranous to fleshy; vernation convolute to conduplicate. Inflorescences terminal or lateral, 1-flowered or racemose to paniculate, 

rarely subumbellate. Flowers small and inconspicuous to large 

and showy, primarily zygomorphic, bisexual (rarely plants monoecious), 3-merous. Sepals 3, free to variously connate, frequently 

1 sepal (dorsal sepal) different from the other 2 (lateral sepals), 

lateral sepals often producing at their bases a spur or mentum 

(i.e., chin-like extension at the base of the flower composed of 

column foot, base of labellum, and lateral sepals). Petals 3, 1 

petal (opposed to the dorsal sepal) modified into a labellum that 

functions as a landing platform for insect visitors, commonly of 

different shape and color than the other 2 petals, often lobed and 

frequently provided with calli, hairs, keels or other ornaments, in 

some cases basally produced into a spur, the other 2 petals similar 

MAP 104. Stromanthe jacquinii occurrence in Venezuela. MAP 105. Stromanthe tonckat occurrence in Venezuela.
NUMBER 100 • 107

FIGURE 18. Marantaceae, Stromanthe tonckat (Aubl.) Eichler. A. Habit (plant in fruit). B. Habit (plant in flower). C, D. Leaf (details showing open sheath, petiole, and pulvinus). E. Flower. F. Fruit with persistent sepals. G. Cross section of fruit showing a seed. (A–D, F, G, Dorr & 

Stergios 8824; E, van der Werff et al. 7487.)
108 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

in shape and color and alternate with the sepals. Stamens and 

styles fused into a complex organ (the column), which is placed 

in the center of the flower. Column with 1 or more, rarely 2 or 

3, fertile stamens located near the apex of the column or on the 

sides; stigmatic surfaces confluent on the ventral face of the column or ± separated, the apex of the medial stigma lobe modified 

to form an organ (the rostellum) that keeps pollen from contacting the stigmatic surface and provides a sticky matter for pollen 

to be attached to pollinators. Anther, when present, placed above 

the rostellum, 2- or 4-locular, located on an apical cavity of the 

column (the clinandrium); pollen in tetrads or more commonly in 

masses (pollinia); pollinia 2, 4, 6 or 8, soft and mealy or hard and 

cartilaginous; pollinia associated with structures that aid in transport and placement over pollinators, the whole complex called 

the pollinarium; pollinarium consisting of the pollinia, a structure 

called viscidium that originates from the rostellum (and hence of 

stigmatic ontogeny) and attaches to the pollinator, caudicles that 

are made of pollinic material (hence originating from the anther), 

and frequently provided with 1 or 2 stipes, which are made of 

anther material and link the pollinia to the viscidium. Ovary inferior, 1–3-locular. Fruit a capsule, dry or rarely fleshy; seeds many, 

very small, dry, wind-dispersed; acotyledonous.

A cosmopolitan, but mainly tropical, family with ~800 genera and 22,000–35,000 species. Approximately 215 genera and 

1,500 species occur in Venezuela.

References. Dunsterville and Garay (1959, 1961, 

1965, 1966, 1972, 1976, 1979a, 1979b, 1979c); Fernández 

(2003); Foldats (1969, 1970a, 1970b, 1970c, 1970d); Pridgeon 

et al. (2001a, 2003, 2005, 2009); Romero-González and Carnevali Fernández-Concha (2000).

The following keys are artificial and do not reflect the relationships of the genera found in Guaramacal. However, subfamily, tribe, and subtribe (if available) are indicated in square 

brackets following each generic description, and these will permit one to discern higher-level relationships. Three subfamilies are found in the park: Epidendroideae, Orchidoideae, and 

Vanilloideae. The vast majority of the genera are placed in the 

Epidendroideae.

KEY TO MAJOR GROUPS OF ORCHIDACEAE

1a. Leaves plicate with (3–)5–9(–15) equally conspicuous nerves or herbaceous (i.e., without conspicuous nerves and grassy 

or herbaceous texture); prefoliation convolute (i.e., leaves rolled during development) . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Inflorescences emerging from the (lateral) base of pseudobulbs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subkey I

2b. Inflorescences emerging from apices of elongate or less commonly short stems and in some terrestrial plants emerging 

from pseudobulbs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subkey II

1b. Leaves conduplicate or terete with midrib more conspicuous than other veins; prefoliation conduplicate (i.e., leaves 

folded along the midrib in development) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Pseudobulbs present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Inflorescences terminal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subkey III

 4b. Inflorescences lateral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subkey IV

3b. Pseudobulbs absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 5a. Inflorescences terminal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subkey V

 5b. Inflorescences lateral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subkey VI

SUBKEY I: LEAVES PLICATE; INFLORESCENCES LATERAL

1a. Pseudobulbs absent; plants terrestrial; leaves distichous; inflorescences terminal and/or lateral . . . . . . . . . Corymborkis

1b. Pseudobulbs present; plants epiphytic or terrestrial; leaves at apices of pseudobulbs or a few leaves on sides of pseudobulbs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Plants terrestrial with homoblastic pseudobulbs (i.e., with 2–5 ± equally developed internodes); inflorescences erect; 

flowers blue or violet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Warreella

2b. Plants epiphytic with heteroblastic pseudobulbs (i.e., with 1 well developed internode); inflorescences pendulous; 

flowers white, yellow or orange or white-spotted with dark purple, never completely violet or blue . . . . . . . . . . . 3

 3a. Leaves 2–4 at the apex of a pseudobulb; inflorescences (3–)6–10(–20)-flowered; flowers resupinate; labellum not 

enclosed by lateral sepals and these not forming a sac . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acineta

 3b. Leaf 1 at the apex of a pseudobulb; inflorescences 1–5-flowered; flowers not resupinate; labellum hidden in the 

sepaline sac . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schlimmia

SUBKEY II: LEAVES PLICATE OR HERBACEOUS; INFLORESCENCES TERMINAL

1a. Stems elongate, conspicuous, erect or rarely prostrate; leaves distichous or partially spiral, distributed along the stem

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
NUMBER 100 • 109

2a. Leaves plicate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Inflorescences branched, terminal and lateral; flowers yellow or creamy yellow; anther much longer than wide, 

located on the back of the column . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Corymborkis

 3b. Inflorescences simple or (seemingly) 1-flowered; flowers white, lilac, purple to orange, never yellow; anther 

shorter than wide, located at the apex of the column . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Inflorescences greatly abbreviated; rachis completely covered by 1 or more spathes forming a conical or 

ellipsoid structure, flowering successive and the inflorescence appearing 1-flowered; flowers large and showy, 

purple or white; petals and sepals >5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sobralia

 4b. Inflorescences elongated or abbreviated, clearly multiflowered; rachis naked or partially covered by bracts 

or spathes; flowers small, showy or not, white, lilac, purple to orange; petals and sepals <2.5 cm long

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elleanthus

2b. Leaves not plicate, herbaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 5a. Inflorescences racemes of 5 or more flowers; flowering ± simultaneous; flowers, usually white or greenish . . . . . 6

 6a. Stems completely erect; 1 or 2 irregularly shaped tubers and fibrous roots emerging from the stem base; 

nectary of the labellum longer than the central lobe of the labellum; petals 2-lobed (split longitudinally into 

2 segments of unequal length); floral segments widely spreading . . . . . . . . . . . . . . . . . . . . . . . . Habenaria

 6b. Stems prostrate or decumbent, erect only in the upper 2⁄3–1⁄2 of their length; fleshy-fibrous roots arising from 

internodes lying on the substrate; nectary of the labellum shorter than the central lobe of the labellum; petals 

entire; floral segments parallel to the column and creating a tubular flower . . . . . . . . . . . . . . . Microchilus

 5b. Flowers solitary or in short racemes of 1–3 flowers; flowering ± successive; flowers green with purple areas on 

the labellum or entirely lilac . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 7a. Understory plants, trailing basally; leaves (in living material) dark green, drying dark brown, almost black, 

short but conspicuously petiolate; flowers green with purple spots on the labellum; floral segments 2–2.5 cm 

long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Psilochilus

 7b. Plants of exposed locations, fully erect; leaves (in living material) light green to glaucous, drying brown, completely sessile and conspicuously sheathed; flowers lilac or purple; floral segments >3 cm long . . . . . . Cleistes

1b. Stems short, inconspicuous or completely to partially underground; leaves in basal rosettes, erect to prostrate . . . . . . 8

8a. Plants with pseudobulbs completely enclosed within a pseudostem (formed by erect petioles and sheathing leaves) 

from which the inflorescence emerges; leaf blades ovate at the apex of the pseudostem and appearing to be sessile, 

seemingly opposite or subopposite; inflorescences umbels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Malaxis

8b. Plants without pseudobulbs or pseudostems; leaf blades elliptic to ovate, rosulate or subdistichous; inflorescences 

spikes or racemes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

 9a. Petals and labellum adnate to column in their basal parts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ponthieva

 9b. Petals and labellum free from column . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

 10a. Inflorescences lateral; leaves often withered and absent at anthesis (i.e., plants hysteranthous) . . . . Pterichis

 10b. Inflorescences terminal; leaves present at anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

 11a. Pedicels conspicuous (longer than floral segments); floral bracts much shorter than the flowers and not 

exceeding the pedicels in length at anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cranichis

 11b. Pedicels inconspicuous (shorter than floral segments); floral bracts exceeding the flowers in length at 

anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

 12a. Dorsal sepal connate at base with lateral sepals to form a tube or a cup . . . . . . . . . Prescottia

 12b. Dorsal sepal completely free from lateral sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

 13a. Column sigmoid, usually pubescent or papillose; leaves basal and cauline, but mostly 

grouped together near base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gomphichis

 13b. Column erect, glabrous; leaves rosulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aa

SUBKEY III: LEAVES CONDUPLICATE; PSEUDOBULBS PRESENT; INFLORESCENCES TERMINAL

1a. Plants vegetatively reduced with leaves <5 cm long; pseudobulbs usually partially hidden by the leaf sheaths; peduncles laterally flattened; column very short, partially covered with warts or rigid black hairs; rostellum elongated into 

a hook; pollinarium of 4 pollinia superimposed on a long, narrowly obtriangular stipe; viscidium linear, hook-shaped

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telipogon

1b. Plants vegetatively variable, but leaves usually >5 cm long; pseudobulbs always conspicuous and never completely covered by the leaf sheaths; peduncles terete or angled, but not laterally flattened; column elongate (at least 1⁄2 the length 
110 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

of the labellum), not covered with hairs or warts; rostellum transverse in the column, inconspicuous, not hook-shaped; 

pollinarium of 4 pollinia with caudicles, without a stipe or hook-shaped viscidium . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Pseudobulbs cylindrical, 8–20× longer than wide, in groups superimposed on top of each other originating from the 

apex of the oldest pseudobulb (when old pseudobulbs flower, flowers can appear lateral); inflorescences with greatly 

reduced rachises and peduncles and flowers solitary or in small, 1–3-flowered fascicles; column with a well-developed 

foot and labellum articulate at the apex of the columnar foot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scaphyglottis

2b. Pseudobulbs fusiform (sometimes appearing subcylindric in the herbarium), <6× longer than wide, not superimposed 

on top of each other; inflorescences with well-developed rachises and peduncles (but peduncles abbreviated in Epidendrum jajense); column without a foot and labellum completely or partially joined to the column . . . . . . . . . . 3

 3a. Labellum free from the column or fused only in its basal 1⁄4; petals wider than sepals; plants and flowers precipitating glycoside crystals in all tissues preserved in alcohol . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prosthechea

 3b. Labellum completely fused to the apex of the column; petals conspicuously narrower than the sepals; plants and 

flowers not precipitating glycoside crystals in all tissues preserved in alcohol . . . . . . . . . . . . . . . . Epidendrum

SUBKEY IV: LEAVES CONDUPLICATE; PSEUDOBULBS PRESENT; INFLORESCENCES LATERAL

1a. Inflorescences 1-flowered, solitary or fascicled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Flowers with a conspicuous spur (nectary projecting toward the back of the flower); pseudobulbs very small and 

almost completely hidden by the sheaths subtending them . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichocentrum

2b. Flowers spurless; pseudobulbs conspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Labellum always articulate at the foot of the column and mobile; petals and sepals >2.5 cm long; petals at least 

4× longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Maxillaria

 3b. Labellum fused rigidly to the column at its base; petals and sepals <2 cm long; petals <3× as long as wide . . . . . 4

 4a. Flowers red or bright yellow (rarely white with a red dot on the labellum), medium-sized; labellum sacshaped or very concave, very fleshy; sepals ~1 cm long; most recent pseudobulbs, at least, with 1 or 2 sheaths 

with foliar blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithidium

 4b. Flowers white, very small; labellum lobed, never deeply concave or extremely fleshy; sepals >0.6 cm long; 

sheaths covering pseudobulbs lacking foliar blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camaridium

1b. Inflorescences few- or many-flowered, but appearing 1-flowered when flowering is successive (persistent pedicels or floral 

bracts provide evidence of the actual few- or many-flowered inflorescence condition) . . . . . . . . . . . . . . . . . . . . . . . . . 5

5a. Pseudobulbs inconspicuous, <1 cm long, often completely or partially covered by sheaths, usually 1-foliolate . . . . . 6

 6a. Plants tiny; leaf blades 2.5–4 × >1 cm; peduncles laterally flattened; flowers without a spur; column very short, 

partially covered with warts; rostellum elongated, hook-shaped; pollinarium with a linear, hook-shaped viscidium 

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telipogon

 6b. Plants larger; leaf blades at least 5 × 1.5 cm; peduncles terete (cylindrical); flowers with a ± conspicuous spur; 

column longer (at least 1

⁄5 the length of the labellar blade) and if verruculose, only in the anther; pollinarium with 

a variable but never elongated, hook-shaped viscidium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 7a. Leaves very fleshy, mottled with purple; spur very conspicuous, significantly longer than blade of the labellum, fully formed by base of the labellum; anther verruculose; flowers showy and fleshy, white or pink with 

red or purple stripes or spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichocentrum

 7b. Leaves coriaceous, less fleshy, uniformly green; spur not very conspicuous, shorter to slightly longer than 

blade of the labellum, created by a basal extension of the lateral lobes within which is inserted basal projections of the labellum; anther smooth; flowers ± showy, membranous to subcoriaceous, red or yellow

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Comparettia

5b. Pseudobulbs conspicuous, ≥2 cm long, often completely or partially uncovered (even though they may be covered by 

conspicuous sheaths), usually 2- or 3-foliate apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

 8a. Pseudobulbs 1-foliate apically; sheaths covering pseudobulbs without foliar blades (or foliar blades much reduced 

and very different from the leaves at the apex of the pseudobulbs); inflorescences racemes . . . . . . . . . . . . . . 9

 9a. Inflorescences pendulous or arching; peduncles ± slender; leaves coriaceous; petals and sepals linear, acute, 

white, creamy or pale pink; labellum blade oblong-subquadrate; labellum base parallel to the column; 

clinandrium fimbriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichopilia

 9b. Inflorescences stiffly erect; peduncles robust (at least 3–4 mm thick); leaves chartaceous-coriaceous; petals 

and sepals broadly elliptic or oblong-elliptic, rounded or truncate, yellowish-brown with darker brown 

areas; labellum blade pandurate (i.e., guitar-shaped); labellum base at an 180° angle to the column; clinandrium smooth or at most erose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Otoglossum
NUMBER 100 • 111

 8b. Pseudobulbs 1- or 2-foliate apically; sheaths covering pseudobulbs with foliar blades very similar (albeit smaller 

at times) to the leaves at the apex of the pseudobulbs; inflorescences panicles or racemes . . . . . . . . . . . . . . 10

 10a. Inflorescences short racemes, shorter to somewhat longer than the leaves at the apex of the pseudobulb; 

floral bracts glumaceous (chartaceous), exceeding the flowers in length; sepals 9–13 mm long; flowers 

tubular, yellow or orange; labellum rigidly fused to the base of the column and forming a saccate nectary 

that is inserted into the receptacle of the flower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brassia

 10b. Inflorescences panicles or racemes, usually longer than the leaves at the apex of the pseudobulb; floral 

bracts never glumaceous (chartaceous) or exceeding the flowers in length; sepals usually >15 mm long 

(~10 mm in Cyrtochilum megalophium); flowers with widely opening parts, variously colored; labellum 

completely free of the column (but not articulate with it), not forming a nectary with the column . . . 11

 11a. Pseudobulbs rounded or ovate, surface dull, often hidden by large, foliaceous, distichous bracts; roots 

relatively thick; pollinia with relatively broad stipe and large caudicles . . . . . . . . . . . . Cyrtochilum

 11b. Pseudobulbs compressed, ancipitous (i.e., distinctly edged), surface glossy; roots relatively thin; 

pollinia with large stipe and short caudicles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

 12a. Basal portion of labellum parallel (and often adnate) to the basal portion of the column for >1⁄2

the length of the column . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Odontoglossum

 12b. Basal portion of labellum spreading from the base of the column, parallel for <1⁄4 the length of 

the column . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oncidium

SUBKEY V: LEAVES CONDUPLICATE OR TERETE; PSEUDOBULBS ABSENT; INFLORESCENCES TERMINAL

1a. Leaves arranged along the stem or when there is a single stem 2 or more leaves at the apex of the stem; pedicels not 

articulate with the ovary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Leaves terete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jacquiniella

2b. Leaves dorsoventrally compressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Epiphytic plants; vegetatively reduced with leaf blades usually <5 cm long; peduncles flattened laterally; column 

very short, partially covered with warts or stiff black hairs; rostellum elongated in the shape of a hook; pollinarium with 4 pollinia superimposed on a long, narrowly obtriangular stipe, viscidium linear, hook-shaped

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telipogon

 3b. Epiphytic or terrestrial plants; vegetatively variable but leaf blades usually >5 cm long; peduncles terete or angled 

but not flattened laterally; column elongated (at least 1⁄2 the labellar length), not covered with warts or hairs; rostellum transverse in the column, inconspicuous, not hooked; pollinarium with 4 pollinia with caudicles, without 

stipe or hook-shaped viscidium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Stems 10–20× longer than wide with 2 or 3 leaves at the apex of a stem; stems superimposed each on top of the 

other; flowers with labellum articulate with the column foot; column foot well developed . . . . Scaphyglottis

 4b. Stems with leaves distributed along length of stem or when leaves rarely concentrated at the apex of the stem, 

the stem much shorter (no more than 5× longer than wide); stems not superimposed each on top of the other; 

flowers with column rigidly fused to labellum its entire length; column foot lacking . . . . . . . Epidendrum

1b. Leaves solitary at the apex of the stem (or stems); pedicels articulate with the ovary (Pleurothallidinae) . . . . . . . . . . . 5

5a. Stems filiform; sheaths lepanthiform (i.e., sheaths inflated and cupuliform, open and spreading apically with ciliate 

margins, the nerves prominent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 6a. Petals wider than long, clearly 2-lobed; labellum with a ± conspicuous, deeply 2-lobed basal claw, lobes attenuate 

basally where they unite to form a labellar claw, sinus of the apical junction of the 2 lobes with a small “appendage” (ontogenetically homologous with the apical lobe) of various shapes, usually pubescent; column ± elongate, 

apically embraced by the lateral lobes of the labellum, ventral faces of the labellum in the same plane as the back 

of the column; anther apical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepanthes

 6b. Petals longer than wide, not lobed; labellum without a claw or claw inconspicuous, simple or 3-lobed, without 

an appendage; column short or ± elongate, not closely embraced by the lateral lobes of the labellum (or labellum 

simple); anther apical or subventral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 7a. Anther subapical on a short column (<1⁄3 the length of the labellum); inflorescences 1-flowered (our species)

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepanthopsis

 7b. Anther ventral or apical on a column that is at least 1⁄2 the length of the labellum; inflorescences 2–6-flowered 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichosalpinx

5b. Stems filiform or more commonly more robust, without lepanthiform sheaths, sheaths tubular, margins entire, not 

ribbed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
112 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

 8a. Sheaths covering stems hispid (covered with black hairs); inflorescences 1-flowered; petal apices thickened in 

black globules (osmophores) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myoxanthus

 8b. Sheaths covering stems not hispid; inflorescences 1- to many-flowered; petal apices not thickened in black 

globules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

 9a. Stems inconspicuous, much shorter than the leaves (1

⁄5 the length of the leaf blade), attenuate in a conspicuous 

pseudopetiole that can be mistaken for a stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

 10a. Sepals completely fused in a tube, apically free and attenuate in conspicuous tails 1.5–4 cm long; petals 

and labellum much smaller than the sepals and completely enclosed in the sepaline tube; petals with a 

retrorse tooth on the inner margin (opposite the column) . . . . . . . . . . . . . . . . . . . . . . . . Masdevallia

 10b. Sepals completely free or, if fused in a basal tube, then the tails <1 cm and the tube <5 mm long; petals 

and sepals variable, usually not completely enclosed within the sepaline tube; petals without teeth on 

the inner margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

 11a. Column wider than long; stigma 2-lobed, the lobes subapical; flowers pink or translucent pink 

with a darker labellum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platystele

 11b. Column conspicuously longer than wide; stigma simple; flowers yellow or greenish translucent

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Specklinia

 9b. Stems conspicuous, at least 1⁄2 the length of the leaf blade, attenuate in a pseudopetiole or not . . . . . . . 12

 12a. Inflorescences 1-flowered, solitary or fascicled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

 13a. Leaf bases cordate; leaf blades broadly ovate to linear; lateral sepals completely fused; pollinia 2 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acronia

 13b. Leaf bases not cordate; leaf blades linear to elliptic; lateral sepals free or fused in the basal 1⁄4; 

pollinia 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pleurothallopsis

 12b. Inflorescences 2- to many-flowered, simultaneous or successive (in which case inflorescences may appear 1-flowered but a careful examination of the rachis will reveal scars left by earlier flowers), solitary 

or multiple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

 14a. Inflorescences associated with an annulus (i.e., an articulation on the stem just to the side of the 

base of the inflorescence below the articulation with the leaf blade), subtended by a ± conspicuous 

spathe emerging from 1 side of the stems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

 15a. Sepals broadly triangular, very shortly fused at the base and widely spreading (in most species) making the flower resemble a triangle; petals transverse and very fleshy, calli often in 

the apical margin; labellum very fleshy and resembling 1⁄8 of a sphere (in 3 dimensions); 

stigma 2-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stelis

 15b. Sepals oblong or ovate, not broadly triangular, the flower not resembling a triangle; petals longer 

than wide, without calli in the apical margin (or calli inconspicuous); labellum longer than 

wide, membranous, oblong, elliptical or ovate in general outline; stigma 1-lobed . . . . . . . . 16

 16a. Flowers relatively large and showy; dorsal sepal 3–6.7 cm long; perianth segments 

translucent yellow with conspicuous wine-red nerves; labellum >10 mm long, with 

hooked and porrect (i.e., extended horizontally) lateral lobes . . . . . . . . . Lindleyalis

 16b. Flowers small or medium-sized; dorsal sepal ≤2 cm long; perianth segments variously 

colored; labellum <8 mm long, with various lateral lobes but not hooked and porrect

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

 17a. Labellum articulate at its base with the globular apex of the columnar foot 

(forming a ball-shaped joint); labellum 3-lobed; sepals fused in their basal 1⁄2

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crocodeilanthe

 17b. Labellum articulate with the column on a transverse strip at the base of the labellum and apex of the column; labellum simple or inconspicuously 3-lobed and 

its basal margins erect or not; sepals free or lateral sepals connate for a variable 

length (often fully connate) as a synsepal . . . . . . . . . . . . . . . . . . . . . Anathallis

 14b. Inflorescences not associated with an annulus; spathes, if present, emerging from leaf bases . . . . 18

 18a. Stems triangular in cross section, winged; flowers very fleshy; inflorescences reclining or 

resting on the leaves; clinandrium laciniate or toothed; anther subventral; stigma ventral

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acianthera

 18b. Stems terete in cross section, not winged; flowers membranous; inflorescences various, often 

pendulous, rarely resting on the leaves; clinandrium entire; anther apical or dorsal; stigma 

apical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pleurothallis
NUMBER 100 • 113

SUBKEY VI: LEAVES CONDUPLICATE; PSEUDOBULBS ABSENT; INFLORESCENCES LATERAL

1a. Leaf blades laterally compressed, not articulate with leaf sheaths, densely imbricate basally (stem completely covered with 

leaves except for the bases of old stems); inflorescences of several successive flowers (sometimes >1 flower present when 

the inflorescence branches) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lockhartia

1b. Leaf blades dorsoventrally compressed, articulate or not with leaf sheaths, loosely or subdensely arranged on the stem; 

inflorescences composed of 1 flower or with 2 flowers in a subumbel of ± simultaneous flowers . . . . . . . . . . . . . . . . . 2

2a. Large terrestrial (or possibly and less commonly epiphytic) plants; leaf blades ≥10 cm long; stems ≥60 cm long; column completely fused with the labellum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epidendrum

2b. Small epiphytic or humicolous plants; leaf blades <3 cm long; stems ≤30 cm long; column free from the labellum . . . . 3

 3a. Inflorescences 1-flowered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Plants pendulous; fruits muricate, suborbicular in cross section; leaves marcescent; labellum attenuate at base 

and anchor-shaped at apex (i.e., apex 2-lobed with retrorse lobes) . . . . . . . . . . . . . . . . . . . . . . . . . Dichaea

 4b. Plants erect; fruit not muricate; leaves not marcescent; labellum not anchor-shaped at apex. . . . Brachionidium

 3b. Inflorescences 1–5-flowered, in a small umbel of ± simultaneous flowers . . . . . . . . . . . . . . . . . . . Fernandezia

Aa Rchb. f.

Aa Rchb. f., Xenia Orch. 1: 18. 1854.

Terrestrial herbs. Roots fleshy, linear-fusiform, fascicled at 

the base of the rosette. Leaves rosulate, 3 or 4 leaves per rosette, present (our species) or absent at anthesis (i.e., plants 

hysteranthous); leaf blades elliptic or linear-elliptic, bases very 

short pseudopetiolate. Inflorescences terminal, originating from 

the remnants of the rosette of the previous growing season and 

thus appearing to be lateral, racemose and appearing spiciform 

because of short pedicels, racemes relatively short compared to 

peduncles, entirely covered by floral bracts; peduncles robust, 

completely surrounded by tubular, ± appressed, imbricate, 

hyaline sheaths; rachises with flowers densely packed, manyflowered; floral bracts hyaline to diaphanous, usually longer than 

flowers, erect, reflexed after anthesis. Flowers not resupinate, 

small, inconspicuous, glabrous; perianth segments membranous. 

Sepals free or sometimes shortly connate at base, ± similar or lateral sepals much larger than dorsal sepal, often recurved. Petals 

free, narrower than sepals, glabrous. Labellum calceolate (i.e., 

slipper-shaped), margins involute, fimbriate or denticulate, with 

a pair of subglobose calli basally; column erect, short, somewhat 

dilated apically, glabrous; clinandrium relatively small, without 

lobes, never tubulose; rostellum transverse, truncate, reniform, 

relatively large; anther dorsal, erect, relatively small; pollinia 4, 

linear-oblong, sessile. Ovary glabrous to glandular-pubescent. 

[Orchidoideae: Cranichideae: Cranichidinae.]

A genus of 25–30 species found at high elevations in South 

America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, 

and Argentina). Two species are known from Venezuela.

References. Álvarez-Molina and Cameron (2009); 

Pridgeon et al. (2003); Salazar Chávez et al. (2009).

The species of Aa are heliophilous and often found in light 

gaps in forest.

Salazar Chávez et al. (2009) determined Aa to be monophyletic and closely related to Porphyrostachys Rchb. f., the two 

genera forming a clade sister to Gomphichis Lindl., which also 

occurs in Guaramacal. Álvarez-Molina and Cameron (2009), 

however, recovered a slightly different topology. These same 

three genera and two or three others formed a highly supported 

clade (Altensteinia clade) of predominantly Andean genera. A 

relationship between Aa and Myrosmodes Rchb. f. was not completely resolved, and Álvarez-Molina and Cameron (2009) suggested that with more taxon sampling Myrosmodes ultimately 

might prove to be better accommodated in Aa. Álvarez-Molina 

and Cameron (2009) also recovered a relationship between Aa/

Myrosmodes, Porphyrostachys, and Gomphichis similar to that 

of Salazar Chávez et al. (2009), who did not sample Myrosmodes.

Aa hartwegii Garay, Fl. Ecuador 9: 161, fig. 49. 1978; Romero 

and Carnevali, Orchids Venez., 2nd ed., 1: 1. 2000; Morillo, 

in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 

92–93, fig. 23. 2011.

Altensteinia paleacea auct., non (Kunth) Kunth; Foldats, Fl. Venez. 15(1): 

403, fig. 151. 1969, pro parte.

MAP 106. Aa hartwegii occurrence in Venezuela.
114 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Terrestrial herbs. Leaf blades 15–18 × 1.6–1.8 cm; pseudopetioles 2–4 cm long. Inflorescences 7–9 cm long; peduncles 

45–48 cm long. Floral bracts ovate-lanceolate, 11–13 × 5 mm, 

acute. Flowers white. Dorsal sepal 2.5 × ~1 mm.

Found in South America (Colombia, Venezuela, and Ecuador). In Venezuela, known only from the Andes (Táchira and 

Trujillo; Map 106). In the park, known from the Laguna del 

Pumar and vicinity; 3,000 m.

Acianthera Scheidw.

Acianthera Scheidw., Allg. Gartenzeitung 10: 292. 1842. Pleurothallis subgen. Acianthera (Scheidw.) Luer, Monogr. Syst. Bot. Missouri Bot. 

Gard. 20: 12. 1986.

Epiphytic, lithophytic or rarely subterrestrial herbs; caespitose or repent, rarely pendent. Rhizomes reduced or creeping. 

Stems terete, laterally compressed or triangular in cross section 

and apparently winged, reduced to much longer than the leaves, 

frequently covered by 1 to several tubular and appressed sheaths. 

Leaves erect, perpendicular or deflexed on the stem; leaf blades 

commonly ovate or ovate-elliptic, more rarely linear, suborbicular to obovate or subcylindrical, bases attenuate, cordate or decurrent on stem, apices slightly 3-toothed, coriaceous or fleshy, 

sessile of petiolate. Inflorescences terminal, not associated with 

an annulus, emerging from the apex of the stem or appearing 

to originate directly from the leaf when leaf margins are decurrent on triquetrous stems, solitary or less commonly fasciculate, 

racemose or 1-flowered by abortion, usually shorter than subtending leaves and frequently appressed to the ventral face of 

same, rarely pendulous, flowers opening successively or more 

commonly ± simultaneously; floral bracts usually tubular, fleshy, 

less commonly large and conspicuous. Flowers commonly resupinate, small or medium-sized, variously colored, often malodorous, fleshy (compared to other Pleurothallidinae). Sepals almost 

free or, more commonly, the lateral sepals connate for a variable 

length (often their full length) in a synsepal that supports the 

labellum, externally smooth or verruculose; dorsal sepal often 

narrower and differently shaped than synsepal. Petals less fleshy 

and conspicuously smaller than sepals, generally parallel to the 

column, margins smooth or frequently dentate to fimbriate or laciniate. Labellum very fleshy, attached to the column foot, simple 

to deeply 3-lobed, lateral lobes relatively small, erect, and clasping the column, apical lobe usually verruculose, margins smooth, 

more commonly erose to laciniate; disk usually with 2 longitudinal calli in the basal or medial portion; column elongate, semicylindrical, straight or ± arcuate, with a well-developed foot, 

frequently glandular or verruculose; clinandrium usually erose 

or dentate to fimbriate; anther operculate, subventral; pollinia 

2; stigma and rostellum ventral. Ovary terete or subtriangular, 

glabrous or pubescent, attached to pedicel. Capsules ellipsoid to 

pyriform. [Epidendroideae: Epidendreae: Pleurothallidinae.]

A neotropical genus of over 250 species found in Mexico, 

Central America, the West Indies, and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Argentina, and 

Uruguay). One center of diversity is in southeastern Brazil. 

Twenty-four species are reported from Venezuela.

References. Luer (1986d, 2002b, 2004); Pridgeon 

and Chase (2001); Pridgeon et al. (2001b, 2005).

Although Luer (1986d, 2002b, 2004) treated Acianthera

as a subgenus in a broadly defined Pleurothallis R. Br., there is 

strong molecular support (Pridgeon et al., 2001b) for recognizing Acianthera as a distinct genus.

KEY TO THE SPECIES OF ACIANTHERA

1a. Leaf blades ovate or ovate-elliptic, 3–5(–8) × 1.5–2.5(–4) cm; stem wings not decurrent on the leaf base; flowers opening 

widely, without side windows (i.e., openings created by the way the concave dorsal sepal rests on the erect apex of the 

synsepal); dorsal sepal oblong–obovate; margins of lateral sepals entire, apices acute or obtuse, inconspicuously apiculate; petals entire or margins slightly erose or serrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. casapensis

1b. Leaf blades elliptic, 6–10 × 2–3 cm; stem wings conspicuously decurrent on the leaf base; flowers globular with side 

windows (see above); dorsal sepal obovate; margins of lateral sepals erose-fimbriate, apices truncate and conspicuously 

apiculate; petals strongly laciniate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. prognatha

Acianthera casapensis (Lindl.) Pridgeon & M. W. Chase, Lindleyana 16: 242. 2001. Pleurothallis casapensis Lindl., Edwards’s Bot. Reg. 28: Misc. 76. 1842; Dorr et al., Contr. 

U.S. Natl. Herb. 40: 52. 2000 [2001].

Pleurothallis chamensis Lindl., Orchid. Linden. 2. 1846; Dunsterville and 

Garay, Venez. Orchids Ill. 1: 314–315. 1959; Foldats, Fl. Venez. 15(2): 

235–238, fig. 269. 1970; Romero and Carnevali, Orchids Venez., 2nd 

ed., 3: 764. 2000; Fernández, Orquídeas Nat. Táchira 176. 2003. Acianthera chamensis (Lindl.) Pridgeon & M. W. Chase, Lindleyana 16: 

242. 2001.

Epiphytic or lithophytic herbs, shortly creeping; stems 3–15 

cm long. Inflorescences 3–10-flowered. Flowers yellowish-green 

to yellow (becoming orange in age). Dorsal sepal 5–9 mm long. 

Labellum simple, 2.4–5.6 mm long.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, Bolivia, and Brazil). In Venezuela, found in the Andes 

(Barinas, Mérida, Portuguesa, Táchira, and Trujillo), the Sierra 

de Perijá (Zulia), the Cordillera de la Costa (Anzoátegui, Aragua, Distrito Federal, Falcón, Miranda, Sucre, and Yaracuy), 

and Nueva Esparta (Map 107). In the park, known from La 
NUMBER 100 • 115

Divisoria de la Concepción and other localities on the south 

slope of Guaramacal; 1,600 m.

Acianthera prognatha (Luer & R. Escobar) Pridgeon & M. W. 

Chase, Lindleyana 16: 245. 2001. Pleurothallis prognatha

Luer & R. Escobar, Orquideología 16: 30, t. p. 29. 1983; 

Dorr et al., Contr. U.S. Natl. Herb. 40: 53. 2000 [2001]; 

Fernández, Orquídeas Nat. Táchira 189. 2003.

Epiphytic herbs, ± caespitose or shortly creeping. Inflorescences 3–6-flowered. Flowers green (our specimens) or dullorange flowers (or only becoming orange in age?). Dorsal sepal 

8.5–9.5 mm long. Labellum obovate, subacute, 7 mm long.

Found in South America (Colombia and Venezuela). In Venezuela, found only in the Andes (Portuguesa and Táchira; Map 

108). In the park, known only from La Divisoria de la Concepción; 1,700 m.

Acineta Lindl.

Acineta Lindl., Edwards’s Bot. Reg. 29: Misc. 67. 1843.

Epiphytic or rarely terrestrial herbs; robust, caespitose. Stem 

modified into ovoid to subcylindrical pseudobulbs arising from 

1 internode, often compressed and/or sulcate. Leaves (1–)2–4, 

articulate; leaf sheaths withering in age; leaf blades narrowly 

ovate, acute, plicate, coriaceous, subpetiolate (petioles indistinct). Inflorescences 1 to several, lateral, many-flowered pendent racemes arising from the base of the pseudobulbs. Flowers 

resupinate, globose, fleshy, waxy, showy; floral bracts concave, 

ovate, shorter than the pedicellate ovary. Sepals widely elliptic to 

very broadly ovate, concave, spreading, fleshy; lateral sepals partially connate or not, asymmetrical. Petals spathulate, generally 

smaller than sepals. Labellum fleshy, rigid, subsessile to conspicuously clawed, 3-lobed, claw concave to subsacciform, lateral 

lobes large, erect, hatchet-shaped, triangular, transversely elliptic 

or subreniform, midlobe subrhombic, flat or apically shallowly 

concave; disk with fleshy calli; column short, erect, semiterete, 

glabrous or slightly pubescent toward the base, apically winged 

to subauriculate, rarely produced into a short, broad foot; anther terminal, operculate, incumbent, 1- or imperfectly 2-locular; 

pollinarium with 2 yellow pollinia, narrowly obovate, cleft, cartilaginous, and dorsoventrally flattened, tegula (i.e., covering) 

long, oblong-linear; viscidium subtriangular or bifurcate; rostellum ligulate. [Epidendroideae: Cymbidieae: Stanhopeinae.]

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, and 

Brazil). Acineta comprises 10−15 species, 3 of which are found 

in mountainous parts of Venezuela.

References. Christenson (2006); Gerlach (1999b); 

Pridgeon et al. (2009); Whitten et al. (2000).

Whitten et al. (2000) identified the monotypic Bolivian 

Vasqueziella Dodson as sister to Acineta and speculated that it 

might, in fact, be synonymous.

Acineta cryptodonta Rchb. f., Bonplandia (Hannover) 2: 92. 

1854; Fernández, Orquídeas Nat. Táchira 31. 2003; Llamozas et al., Libro Rojo Fl. Venez. 380. 2003.

Acineta erythroxantha auct., non Rchb. f.; Foldats, Fl. Venez. 15(4): 185–

187, fig. 638. 1970.

Acineta sella-turcica auct., non Rchb. f.; Foldats, Orquideophilo 3(1): 14. 

1995.

MAP 107. Acianthera casapensis occurrence in Venezuela.

MAP 108. Acianthera prognatha occurrence in Venezuela.
116 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Epiphytic herbs, 30–50 cm tall; pseudobulbs 8–10 cm tall, 

fusiform to pyriform, apically 2- or 3-foliate. Leaf blades 25–60 

cm long, pendulous. Flowers yellow. Sepals 3.5–7 cm long.

Found in South America (Colombia and Venezuela). In Venezuela, found only in the Andes (Barinas, Lara, Táchira, and Trujillo; Map 109). The presence of this species in the park is not 

vouchered by an herbarium specimen; we include it on the basis 

of a sight record by G. Gerlach (pers. comm.). Gerlach did not indicate the elevation in the park where he encountered this species, 

but other Venezuelan collections have been made at 1,800 m.

Acronia C. Presl

Acronia C. Presl, Rel. Haenk. 1: 103. 1827.

Zosterophyllanthos Szlach. & Marg., Polish Bot. J. 46: 118. 2001.

Epiphytic, lithophytic or subterrestrial herbs; caespitose to 

repent, erect to pendent. Rhizomes very short to elongate. Roots 

few to many, slender to coarse and fleshy. Stems consisting of 

modules with a short basal part and an aerial part of 2 or 3 internodes, with 1 apical leaf, internodes covered by tubular, appressed sheaths, sheaths shorter than internodes, aerial modules 

simple or, more rarely, proliferous (becoming scandent), usually longer than the leaves, terete or rarely laterally compressed. 

Leaves attached at apex of stem; leaf blades usually ovate and 

bases cordate or subcordate (leaf blades deflexed in some species 

with cordate bases), less commonly elliptic, linear or lanceolate 

and bases cuneate to pseudopetiolate, apices obtuse, acute to 

abruptly short-acuminate with a small mucro in the center of the 

apical sinus, membranous to coriaceous. Inflorescences solitary 

(rarely fasciculate), 1- or few-flowered racemes, almost always 

shorter than leaves; peduncles thin or thick, always emerging 

from a small to large and subfoliaceous spathe; floral bracts tubular, usually shorter than floral pedicels. Flowers resupinate or not, 

arising from leaf bases, usually small, sometimes relatively large 

and conspicuous, color variable but most often green, yellow or 

deep burgundy. Sepals with or without keels, glabrous or more 

rarely verruculose or even somewhat pubescent, membranous or 

fleshy; dorsal sepal free; lateral sepals completely connate to form 

a ± concave synsepal that rests on the labellum, synsepal usually 

ovate or ovate-elliptic, obtuse, acute or acuminate, base sometimes with a mentum. Petals free, usually narrower than the sepals, margins entire, papillose or denticulate, straight or recurved, 

narrowly ovate, broadly ovate, lanceolate to obovate, obtuse, 

acute or acuminate, membranous or fleshy. Labellum membranous or fleshy, entire, usually ovate or lanceolate, subquadrate or 

obovate, flat, concave or convex, usually with a well-developed 

glenion (i.e., small area with a shiny surface that appears wet, 

which is on the upper plane of the labellum, where it rests beneath 

the column) at the base, margins and surfaces smooth, calli papillose or margins fimbriate, articulate to rigidly joined to columnar 

base; column semiterete or conical, short, thick, sometimes with a 

small foot, without wings; anther apical; pollinia 2, narrowly pyriform with small caudicles; stigma apical, transverse or 2-lobed. 

Ovary 3-valved, pedicellate, smooth or (more rarely) verruculose, 

deciduous. [Epidendroideae: Epidendreae: Pleurothallidinae.]

A neotropical genus of ~220 species found in Central America and South America (Colombia, Venezuela, Ecuador, Peru, 

Bolivia, Brazil, and Paraguay). Approximately 35 species occur 

in Venezuela.

References. Luer (2005); Wilson et al. (2011).

Acronia, a segregate of Pleurothallis, consists of two sections, sect. Acronia, with inflorescences composed of various 

numbers of flowers, and sect. Macrophyllae-fasciculatae (Lindl.) 

Luer, with 1-flowered inflorescences. Our species belong to this 

latter section.

The generic concept presented here is strictly morphological. 

There is a suggestion (Wilson et al., 2011) based on unpublished 

and preliminary molecular data that if Acronia and Zosterophyllanthos Szlach. & Marg. are shown to be monophyletic, then the 

group would be relegated to subgeneric status in an expanded 

concept of Pleurothallis.

KEY TO THE SPECIES OF ACRONIA

1a. Leaf blades linear, >15× longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. calamifolia

1b. Leaf blades elliptic to broadly ovate, never more than 8× longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Floral pedicels >4 cm long; flowers pendulous or obscured beneath the leaf blade . . . . . . . . . . . . . . . A. ruberrima

2b. Floral pedicels <2 cm long (usually <1 cm long); flowers borne on the upper surface of the leaf blade . . . . . . . . . 3

MAP 109. Acineta cryptodonta occurrence in Venezuela.
NUMBER 100 • 117

 3a. Adult leaf blades <6.5 cm long, usually <2.5 cm wide; adult stems 5–8(–9) cm long . . . . . . . . . . . . . . . . . . . 4

 4a. Dorsal sepal and synsepal ± the same width; labellum rounded and thickened apically; flowers greenishsalmon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sp. A

 4b. Synsepal conspicuously wider than dorsal sepal; labellum acute apically; flowers yellow or yellowish-brown 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. phyllocardioides

 3b. Adult leaf blades >8 cm long (if somewhat shorter, then no more than 2.5× longer than wide), usually >3 cm 

wide; adult stems 9–20 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 5a. Petals obovate or oblong-obovate, 3-nerved; flowers completely yellow; leaf blades oblong-elliptic to ovateelliptic, always 3–4× longer than wide, bases rounded to deeply cordate; labellum broadly oblong . . . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. semiscabra

 5b. Petals at most narrowly oblong, 1-nerved; flowers usually purplish or brownish-purple, sepals sometimes 

translucent yellow or the flower completely yellow but then leaf blades 1.3–2× longer than wide; leaf blades 

ovate or ovate-elliptic, bases cordate to deeply cordate; labellum ovate or ovate-triangular to elliptic . . . 6

 6a. Dorsal sepal (when hydrated) at least 20 mm long (15–17 mm long in herbarium material), 3–5-nerved 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 7a. Labellum elliptic in general outline, margins in the apical 1⁄2 plane; upper 1⁄2 of petals glabrous

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. bivalvis

 7b. Labellum ovate in general outline, margins in the apical 1⁄2 incurved, rolled down so that the labellum 

apex is conical-cylindric; upper 1⁄2 of petals pubescent . . . . . . . . . . . . . . . . . . . . . . . A. siphoglossa

 6b. Dorsal sepal (when hydrated) 8–12 mm long, 3-nerved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

 8a. Flowers purplish-brown; dorsal sepal and synsepal 1.2–1.8× longer than wide; petals oblong, obtuse, 

4–5× longer than wide, margins erose; labellum ovate, apex obtuse or rounded; glenion (see generic description) conspicuous, ovate, covering at least 1⁄3 of the total area of the labellum . . . . A. archidiaconi

 8b. Flowers greenish-yellow translucent, yellowish or pink; dorsal sepal and synsepal at least 3× longer 

than wide; petals linear-triangular, acute, at least 7× longer than wide, margins entire; labellum narrowly triangular, apex acute; glenion (see generic description) inconspicuous . . . . . A. coriacardia

Acronia archidiaconi (Ames) Carnevali & G. A. Romero, comb. 

nov. Pleurothallis archidiaconi Ames, Sched. Orchid. 9: 29. 

1925; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 

751. 2000; Dorr et al., Contr. U.S. Natl. Herb. 40: 52. 2000 

[2001]; Carnevali and Ramírez-Morillo, in Berry et al., Fl. 

Venez. Guayana 7: 512. 2003.

Pleurothallis monocardia auct., non Rchb. f.; Dunsterville and Garay, Venez. 

Orchids Ill. 1: 336–337. 1959; Foldats, Fl. Venez. 15(2): 345–347, fig. 

312. 1970.

Epiphytic herbs, to 25 cm tall, usually shorter. Leaf blades 

7–10 × 3.5–5 cm, bases cordate, apices shortly acuminate, minutely 3-toothed. Flowers purple-brown. Dorsal sepal 5–7 mm 

long. Labellum reddish-purple with a yellow center.

Found in South America (Venezuela, Trinidad and Tobago, 

the Guianas, and Ecuador). Venezuelan collections are from the 

Andes (Portuguesa, Táchira, and Trujillo), the Cordillera de la 

Costa (Aragua, Distrito Federal, Falcón, and Miranda), and 

the Venezuelan Guayana (Amazonas and Bolívar; Map 110). 

We have a single record of this species from the south slope of 

Guaramacal that consists of a photograph taken by B. Stergios 

(see Carnevali and Ramírez in Cuello A., 1999: 93).

Luer (2005: 93–96) includes this species in his concept of 

Acronia bivalvis (Lindl.) Luer but acknowledges the latter is “the 

greatest catch all or waste basket of the genus.”

Pleurothallis omoglossa Luer, described from Ecuador, 

might be conspecific with Acronia archidiaconi. If so, the latter 

name has priority.

Acronia bivalvis (Lindl.) Luer, Monogr. Syst. Bot. Missouri Bot. 

Gard. 103: 93. 2005. Pleurothallis bivalvis Lindl., Orchid. 

Linden. 2. 1846; Dunsterville and Garay, Venez. Orchids 

MAP 110. Acronia archidiaconi occurrence in Venezuela.
118 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Ill. 4: 206–207. 1966; Foldats, Fl. Venez. 15(2): 215–217, 

fig. 259. 1970; Romero and Carnevali, Orchids Venez., 2nd 

ed., 3: 758–759. 2000; Dorr et al., Contr. U.S. Natl. Herb. 

40: 52. 2000 [2001]; Fernández, Orquídeas Nat. Táchira 

175. 2003.

Pleurothallis coriacardia auct., non Rchb. f.; Carnevali and Ramírez, in 

Cuello, Parque Nac. Guaramacal 91, fig. 1999 (“Foto: A. Licata”).

Epiphytic herbs, to 40 cm tall. Adult leaf blades 7–13 × 

3–5(–7) cm, bases cordate to deeply cordate, apices acute to subacute. Flowers relatively large for the complex of species with 

cordate leaves. Dorsal sepal 15–22 cm long, pale pink (at base) 

or yellow with purple nerves; synsepal pale purple with darker 

nerves; petals and labellum dark purple; column pale yellow.

Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, recorded from the Andes (Lara, Mérida, 

Portuguesa, Táchira, and Trujillo), the Sierra de Perijá (Zulia), 

and the Cordillera de la Costa (Distrito Federal and Miranda; 

Map 111). In the park, found in cloud forest on both slopes of 

Guaramacal; 1,800–2,750 m.

Acronia calamifolia (Luer & R. Escobar) Luer, Monogr. Syst. 

Bot. Missouri Bot. Gard. 103: 99. 2005. Pleurothallis 

calamifolia Luer & R. Escobar, Orquideología 20: 40. 1996.

Pleurothallis subtilis auct., non C. Schweinf.; Foldats, Fl. Venez. 15(2): 422–

426, fig. 347. 1970, pro parte; Ortega et al., BioLlania 5: 49. 1987; 

Dorr et al., Contr. U.S. Natl. Herb. 40: 53. 2000 [2001].

Epiphytic herbs, 25–45 cm tall. Leaves erect or becoming 

deflexed on the stem; leaf blades linear, 12–16 × 0.7–0.9 cm, 

bases rounded or subcordate, apices acute. Flowers yellowish. 

Dorsal sepal 2.6–2.8 mm long. Labellum darker yellow.

Found in South America (Colombia and Venezuela). In Venezuela, found only in the Cordillera de Mérida (Portuguesa and 

Trujillo; Map 112). In the park, known from La Divisoria de la 

Concepción, Casa Vicuyal, and along the Boconó–Guaramacal 

road; 1,700–2,300 m.

This is the first report of Acronia calamifolia occurring in 

Venezuela. It is a distinctive species closely related to A. subtilis

(C. Schweinf.) Luer, with which it has been confused in the past. 

The latter species has a proportionately smaller convex labellum and longer and narrower petals. Material collected by G. C. 

K. Dunsterville in La Carbonera (Mérida state) and illustrated 

by him under the name Pleurothallis subtilis C. Schweinf. (see 

Dunsterville and Garay, 1966: 244–245; Romero-González and 

Carnevali Fernández-Concha, 2000: 853) has a relatively small, 

strongly convex labellum with a strongly corrugated surface that 

is reddish maroon. This material probably represents a distinct 

species that has yet to be described.

Acronia coriacardia (Rchb. f.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 103: 117. 2005. Pleurothallis coriacardia

Rchb. f., Bonplandia (Hannover) 2: 26. 1854; Romero and 

Carnevali, Orchids Venez., 2nd ed., 3: 767. 2000; Dorr et 

al., Contr. U.S. Natl. Herb. 40: 52. 2000 [2001]; Carnevali 

and Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 7: 

513–514. 2003; Fernández, Orquídeas Nat. Táchira 177. 

2003; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 246–247, fig. 92. 2011.

Pleurothallis linguifera auct., non Lindl.; Foldats, Fl. Venez. 15(2): 329, 331–

332, fig. 305. 1970.

Pleurothallis lansbergii auct., non Regel; Dunsterville and Garay, Orchids 

Venez., ed. 1, 3: 783. 1979.

MAP 111. Acronia bivalvis occurrence in Venezuela. MAP 112. Acronia calamifolia occurrence in Venezuela.
NUMBER 100 • 119

Epiphytic or sometimes lithophytic herbs, erect, to 35 cm 

tall. Leaves erect or deflexed on the stem; leaf blades ovate to 

broadly ovate, 8–12 × 0.3–0.5 cm, bases cordate, apices acuminate. Flowers greenish-yellow translucent, yellowish or pinkishgreen. Dorsal sepal 9–12 mm long.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). In Venezuela, recorded from Amazonas, Aragua, Bolívar, Distrito Federal, Mérida, Miranda, Táchira, Trujillo, and Yaracuy (Map 113). Montane forest below Páramo del 

Pumar and along the southwestern border of the park; 1,900–

2,200(–2,600) m.

Material from the Venezuelan Guayana identified as Acronia coriacardia shows differences in flower color and size and 

may represent a distinct species.

Acronia phyllocardioides (Schltr.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 103: 174. 2005. Pleurothallis phyllocardioides Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19: 193. 

1923; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 

826. 2000; Fernández, Orquídeas Nat. Táchira 188. 2003. 

Zosterophyllanthos phyllocardioides (Schltr.) Szlach. & 

Marg., Polish Bot. J. 46: 120. 2001. Zosterophyllanthos 

phyllocardioides (Schltr.) Szlach. & Kulak, Richardiana 

6(4): 192. 2006, isonym.

Epiphytic herbs, 12–15 cm tall when mature. Leaf blades 

4.5–6 × 1.8–2.5 cm, bases deeply cordate, apices acute. Flowers 

yellow. Dorsal sepal 4.8–5.5 mm long.

Found in Central America and South America (Colombia, 

Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, known 

only from the Andes (Táchira and Trujillo; Map 114). Collected 

near the village of Guaramacal; ~1,500 m.

Acronia ruberrima (Lindl.) Luer, Monogr. Syst. Bot. Missouri 

Bot. Gard. 103: 184. 2005. Pleurothallis ruberrima Lindl., 

Orchid. Linden. 1. 1846; Dunsterville and Garay, Venez. 

Orchids Ill. 1: 344–345. 1959; Foldats, Fl. Venez. 15(2): 

389–391, fig. 332. 1970; Romero and Carnevali, Orchids 

Venez., 2nd ed., 3: 836. 2000; Fernández, Orquídeas Nat. 

Táchira 191. 2003.

MAP 113. Acronia coriacardia occurrence in Venezuela. MAP 114. Acronia phyllocardioides occurrence in Venezuela.

MAP 115. Acronia ruberrima occurrence in Venezuela.
120 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Epiphytic herbs, 35–60 cm tall. Leaf blades narrowly 

oblong-elliptic, 14–16 × 2.5–3 cm, bases shallowly cordate, apices acuminate. Flowers greenish-pink. Dorsal sepal 17–28 mm 

long. Labellum pink or dark purple with darker nerves.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). In Venezuela, known from Apure, Aragua, Distrito Federal, Mérida, Miranda, Táchira, Trujillo, and 

Zulia (Map 115). In the park, collected at La Punta in sector El 

Santuario; ~1,850 m.

Acronia semiscabra (Lindl.) Luer, Monogr. Syst. Bot. Missouri 

Bot. Gard. 103: 190. 2005. Pleurothallis semiscabra Lindl., 

Orchid. Linden. 2. 1846; Dunsterville and Garay, Venez. 

Orchids Ill. 3: 258–259. 1965; Foldats, Fl. Venez. 15(2): 

409–411, fig. 341. 1970; Ortega et al., BioLlania 5: 49. 

1987; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 

844. 2000. Maxillaria semiscabra (Lindl.) P. Ortiz, Orquideología 18: 101. 1991; Dorr et al., Contr. U.S. Natl. Herb. 

40: 51. 2000 [2001].

Epiphytic herbs, 20–35 cm tall. Leaf blades narrowly 

ovate-elliptic to elliptic, (5.5–)7–10 × (2–)2.5–3.5(–4) cm, bases 

rounded to deeply cordate, apices acuminate. Flowers completely 

yellow. Dorsal sepal 5–9 mm long.

Endemic to Venezuela, where it is found in the Andes (Lara, 

Mérida, and Trujillo) and in the Cordillera de la Costa (Miranda; 

Map 116). In the park, found in montane forest and cloud forest 

on both slopes of Guaramacal; 1,800–2,600 m.

Acronia siphoglossa (Luer & R. Escobar) Luer, Monogr. Syst. 

Bot. Missouri Bot. Gard. 103: 192. 2005. Pleurothallis siphoglossa Luer & R. Escobar, Orquideología 20: 82, tt. pp. 

83, 89. 1996; Romero and Carnevali, Orchids Venez., 2nd 

ed., 3: 848. 2000; Dorr et al., Contr. U.S. Natl. Herb. 40: 

53. 2000 [2001].

Pleurothallis grandiflora auct., non Lindl.; Dunsterville and Garay, Venez. 

Orchids Ill. 4: 218–219. 1966.

Epiphytic herbs, 25–30 cm tall. Leaf blades ovate, 7–10 × 

3.5–5 cm, bases cordate to deeply cordate, apices acuminate. 

Flowers yellowish or translucent; labellum and petals purple. 

Dorsal sepal 18–23 mm long.

Found in South America (Colombia, Venezuela, and Ecuador). In Venezuela, known only from the Cordillera de Mérida 

(Mérida and Trujillo; Map 117). In the park, we have a single 

record of this species growing on the south slope of Guaramacal 

that consists of a photograph taken by A. Licata (Carnevali and 

Ramírez in Cuello A., 1999: 90).

Acronia sp. A

Epiphytic herbs, 10–14 cm tall. Leaf blades ovate-elliptic, 

4–4.5 × 1.6–1.8 cm, bases cordate, apices shortly acuminate. 

Flowers spreading, salmon-orange (in sched.). Dorsal sepal 

5.2 × 3 mm.

Endemic to Venezuela (Trujillo; Map 118). In the park, collected on the north slope of Guaramacal along the trail from the 

Laguna de Aguas Negras to Qda. Salvaje; 1,850–2,100 m.

This undescribed species is similar to the widely distributed 

Acronia phyllocardioides, but it has a wider dorsal sepal and a 

rounded and apically thickened labellum. Vegetatively, the new 

species seems to belong to the A. tridentata (Klotzsch) Luer complex, but in this species complex the species have a labellum that 

is strongly geniculate basally and proportionally wider.

MAP 117. Acronia siphoglossa occurrence in Venezuela.

MAP 116. Acronia semiscabra occurrence in Venezuela.
NUMBER 100 • 121

Anathallis Barb. Rodr.

Anathallis Barb. Rodr., Gen. Spec. Orchid. 1: 23. 1877.

Pleurothallis subgen. Acuminatia Luer, Monogr. Syst. Bot. Missouri Bot. 

Gard. 76: 98. 1999.

Epiphytic, lithophytic or rarely subterrestrial herbs; erect or 

pendulous. Rhizomes short or creeping. Stems terete or ventrally 

canaliculate, greatly abbreviated to much longer than the leaves but 

more commonly subsimilar in length, frequently covered by 1 to 

several appressed, tubular sheaths. Leaves erect to perpendicular 

to the stem; leaf blades commonly elliptic to linear or suborbicular to obovate, rarely subcylindrical, bases usually attenuate, apices 

minutely 3-toothed, coriaceous or fleshy. Inflorescences terminal, 

associated with an annulus and emerging just above it and below 

the foliar attachment, solitary or fascicled, racemose or 1-flowered 

by abortion, with successive or ± simultaneous flowers; floral bracts 

usually reduced, less commonly large and conspicuous, membranous. Flowers commonly resupinate, small or medium, variously 

colored, membranous or subfleshy (compared to other Pleurothallidinae). Sepals almost free or lateral sepals connate for a variable 

length (often fully connate) as a synsepal on which the labellum 

rests, externally smooth or verruculose; dorsal sepal frequently narrower and differently shaped than synsepal. Petals less fleshy than 

the sepals and conspicuously smaller, variable in shape, usually parallel to the column, margins entire or often dentate to fimbriate or 

laciniate. Labellum attached to column foot and often with a pair 

of minute lobes or retrorse teeth at the base on each side of the attachment, lamina deeply 3-lobed to more commonly simple, lateral 

lobes relatively small, erect, apical lobe smooth, verruculose or pubescent, margins smooth to more commonly erose to laciniate; disk 

usually with 2 longitudinal calli basally or medially; column elongate, hemicylindrical and concave ventrally, straight or ± curved, 

often membranous and easily pliable, with a well-developed foot; 

clinandrium usually erose or dentate to fimbriate; anther operculate, 

subventral; pollinia 2; stigma and rostellum ventral. Ovary attached 

to pedicel, terete or subtriangular, glabrous or papillose; pedicel 

usually fleshy, scarcely differentiated from the ovary. Capsules ellipsoid to obovoid. [Epidendroideae: Epidendreae: Pleurothallidinae.]

A neotropical genus of 300–400 species found in Mexico, 

Central America, the Greater Antilles, and South America (Colombia, Venezuela, Trinidad and Tobago, the Guianas, Ecuador, 

Peru, Bolivia, Brazil, and Argentina). The genus is most diverse 

in the Andes; 35–40 species are recorded from Venezuela.

References. Chiron et al. (2012); Karremans et al. 

(2013); Luer (1986d, 1999, 2002b); Pridgeon and Chase (2001); 

Pridgeon et al. (2005).

Morphologically, Anathallis is difficult to circumscribe because it has no obvious synapomorphies. It is easily confused with 

Specklinia Lindl.; both genera have an annulus subtending the inflorescence, which is usually many-flowered with ± membranous 

flowers. Also, both genera are highly variable vegetatively and florally. Nonetheless, species of Anathallis usually have stems longer 

than those of Specklinia, which typically has stems shorter that the 

leaf blades. Also, species of Specklinia typically have pedicels that 

are conspicuously thinner and longer than the ovary.

Karremans et al. (2013) provided molecular evidence that 

suggests Anathallis as defined by Pridgeon and Chase (2001) is 

not monophyletic. They found that A. anderssonii (Luer) Pridgeon & M. W. Chase, A. dolichopus (Schltr.) Pridgeon & M. W. 

Chase, A. rubens (Lindl.) Pridgeon & M. W. Chase, and A. sclerophylla (Lindl.) Pridgeon & M. W. Chase, which they considered 

to be close relatives, did not group together into a monophyletic 

clade and appeared to be interrelated with species of Physothallis Garay and Crocodeilanthe Rchb. f. & Warsz. Chiron et al. 

(2012) examined two of these four species of Anathallis and 

found somewhat different relationships; A. sclerophylla was embedded within a well-supported Anathallis clade, whereas A. rubens was part of a Stelis clade, and the species was transferred to 

Stelis Sw. as S. neorubens Chiron, nom. nov.

KEY TO THE SPECIES OF ANATHALLIS

1a. Leaf blades 8–25 mm wide, at least 4× longer than wide, ± as long as the stems they subtend or longer; sepals completely 

free, lanceolate or linear-lanceolate, apices long-acuminate; petal apices acute or rounded; labellum obovate . . . . . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. acuminata

1b. Leaf blades 15–30 mm wide, ~3× longer than wide, much shorter than the stems they subtend; sepals briefly connate basally, 

linear to oblong, apices acute; petal apices rounded and somewhat apiculate; labellum oblong . . . . . . . . . A. sclerophylla

MAP 118. Acronia sp. A occurrence in Venezuela.
122 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Anathallis acuminata (Kunth) Pridgeon & M. W. Chase, Lindleyana 16: 247. 2001. Dendrobium acuminatum Kunth, in 

H. B. K., Nov. Gen. Sp. [fol.] 1: 286. 1815 [1816]; ibid. 

[qu.] 1: 357. 1815 [1816]. Pleurothallis acuminata (Kunth) 

Lindl., Edwards’s Bot. Reg. 28: Misc. 70. 1842; Dunsterville 

and Garay, Venez. Orchids Ill. 4: 200–201. 1966; Foldats, 

Fl. Venez. 15(2): 197–200, fig. 252. 1970; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 747. 2000; Dorr et al., 

Contr. U.S. Natl. Herb. 40: 52. 2000 [2001]; Fernández, 

Orquídeas Nat. Táchira 173. 2003.

Epiphytic herbs, to 25 cm tall (including the inflorescences). 

Leaf blades 4–7.5(–11) cm long. Inflorescences 10–20 cm long. 

Flowers greenish-yellow. Sepals 15–25 mm long.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). Venezuelan material is from the Andes (Lara, 

Mérida, Táchira, and Trujillo) and the Cordillera de la Costa 

(Distrito Federal and Miranda; Map 119). Collected in montane 

forest on the north slope of the park; 1,800–2,550 m.

Anathallis sclerophylla (Lindl.) Pridgeon & M. W. Chase, Lindleyana 

16: 250. 2001. Pleurothallis sclerophylla Lindl., Edwards’s Bot. 

Reg. 21: t. 1797. 1836 [1835]; Dunsterville and Garay, Venez. 

Orchids Ill. 2: 300–301. 1961; Foldats, Fl. Venez. 15(2): 403–

406, fig. 339. 1970; Ortega et al., BioLlania 5: 49. 1987; Romero 

and Carnevali, Orchids Venez., 2nd ed., 3: 843. 2000; Dorr et 

al., Contr. U.S. Natl. Herb. 40: 53. 2000 [2001]; Carnevali and 

Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 7: 521, fig. 

473. 2003; Fernández, Orquídeas Nat. Táchira 192. 2003.

FIGURE 25H

Epiphytic herbs, to 35 cm tall (including the inflorescences). 

Leaf blades (5–)9–12 cm long. Inflorescences 15–27 cm long. 

Flowers yellow; sepals 7.5–21 mm long.

Found in South America (Colombia, Venezuela, Guyana, 

French Guiana, Ecuador, Peru, Bolivia, and Brazil). Venezuelan 

records are from the Andes (Lara, Mérida, Táchira, and Trujillo), the Cordillera de la Costa (Aragua, Distrito Federal, and 

Yaracuy), the Venezuelan Guayana (Amazonas and Bolívar), and 

Nueva Esparta (Map 120). In the park, collected in forest on the 

north slope of Guaramacal; 1,800–2,350 m.

The generic placement of this species needs to be reevaluated 

since molecular studies have produced two different, contradictory hypotheses; either this species is embedded in an Anathallis

clade (Chiron et al., 2012), or it belongs in a Stelis clade (Karremans et al., 2013).

Brachionidium Lindl.

Brachionidium Lindl., Fol. Orchid. 8: Brachionidium 1. 1859.

Yolanda Hoehne, Arq. Mus. Nac. Rio de Janeiro 22: 72. 1919. Brachionidium sect. Yolanda (Hoehne) Pabst, Bradea 1: 269. 1972.

Epiphytic, humicolous or subterrestrial herbs; caespitose 

to long-creeping; erect or prostrate to climbing or sometimes 

pendulous, occasionally tuft forming. Rhizomes erect or creeping, simple or branched; internodes often abbreviated or more 

commonly elongated between the stems, covered with a few 

tubular sheaths, glabrous or verruculose, sheaths usually mucronate. Roots few to many originating at the base of rhizomes 

(in many of the erect species) or along the internodes in trailing 

plants and in some of the erect ones (where these are decurrent 

from the rhizome to the substrate), thin and somewhat fibrous 

to thick and subfleshy. Stems ascending to erect, always shorter 

than subtending leaves, 1-foliate, usually covered by 2 sheaths 

similar to those of the rhizomes. Inflorescences 1-flowered, 

not associated with an annulus originating semilaterally near 

MAP 119. Anathallis acuminata occurrence in Venezuela. MAP 120. Anathallis sclerophylla occurrence in Venezuela.
NUMBER 100 • 123

the apex of the stem (i.e., the point of abscission between leaf 

and stem). Leaves erect on the stem; leaf blades narrowly to 

broadly elliptic or obovate to broadly spathulate, bases cuneate to narrowly cuneate, apices acute to obtuse, with a slit in 

the apex and mucro in the sinus, smooth to rough or verruculose, various shades of green, sometimes tinged with purple, 

1–3-veined on each side of midrib, submembranous to thickly 

coriaceous, shortly pseudopetiolate. Flowers usually not resupinate, solitary, large and relatively showy compared to the size 

of the plant, usually opening widely, usually ephemeral (lasting 

1 day); peduncles variable in length, much shorter than (our 

species) to much longer than subtending leaves; floral bracts 

cucullate, inflated, acuminate, clasping the abbreviated pedicel 

and often most of the ovary; pedicels robust; perianth segments 

green or yellow suffused with red or purple, or completely 

brown, red or purple (our species), often margins ciliate, membranous to subfleshy. Dorsal sepal free, usually elliptic, rarely 

obovate, apices often acuminate and terminating in a slender 

tail; lateral sepals free, connate in a flat or concave synsepal, 

usually ± similar to dorsal sepal, acute or obtuse, apices often 

acuminate and terminating in a thin tail, often bifid. Petals ± 

similar to dorsal sepal (giving the appearance of a flower with 

4 segments), often slightly oblique, margins often minutely ciliate, apices often acuminate and terminating in a slender tail. 

Labellum articulate at the apex of the column foot, conspicuously smaller than other perianth segments, very fleshy, usually transverse, apex acute, obtuse to rounded, margins smooth 

to fimbriate, sometimes bimarginate (because of the thick and 

fleshy labellum), with or without an apiculus, disk ± carinate 

or embedded into the anterior portion, with an elevated or ± 

flat callus, oblong or ovoid, ± pubescent; callus situated in the 

center of the labellum just above the base and fitting just below 

the column in natural position; column relatively short, robust, 

anterior margin 2-dentate or entire, base of the column forming 

a strong foot at the apex of the ovary; anther apical; pollinia 6 

or 8, unequal in size and in 2 groups or packages, clavate, free 

or attached to a small viscidium; rostellum transverse; stigma 

apical, transverse, 2-lobed, sometimes appearing to be 2 stigmas but actually continuous below the rostellum with receptive surfaces protruding on each side. Ovary smooth, 3-valved. 

Capsules oblong to oblong-obovoid, obtuse, trigonal. [Epidendroideae: Epidendreae: Pleurothallidinae.]

A neotropical genus of 65–70 species found in Central 

America, the West Indies, and South America (Colombia, Venezuela, Guyana, Ecuador, Peru, Bolivia, and Brazil). Nine species 

have been found in Venezuela.

References. Luer (1986d, 1995); Pridgeon et al. 

(2001b, 2005).

Brachionidium tuberculatum Lindl., Fol. Orchid. 8: Brachionidium 1. 1859; Foldats, Fl. Venez. 15(2): 11–13, fig. 189. 1970; 

Fernández, Orquídeas Nat. Táchira 37. 2003; Romero and 

Carnevali, Orchids Venez., 2nd ed., 1: 36. 2000; Morillo, 

in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 

97–98, fig. 4. 2011.

Brachionidium steyermarkii Foldats, Acta Bot. Venez. 3: 306, fig. 1. 1968; 

Foldats, Fl. Venez. 15(2): 10–11. 1970.

Epiphytic or humicolous herbs, erect, to 22 cm tall. Rhizomes robust, rigid, ± erect. Roots at the base. Leaf blades elliptic to broadly elliptic, 10–15 × 7–10 mm. Flowers purple (our 

material) or yellow. Dorsal sepal 7–8 × 5 mm; lateral sepals 

6–7 × 5 mm. Petals 6–7.5 × 3.5–5 mm. Labellum 2.4–2.6 × 3.4–

3.6 mm.

Restricted to South America (Colombia, Venezuela, Ecuador, Bolivia, and Brazil). In Venezuela, known only from the 

Andes (Mérida, Táchira, and Trujillo; Map 121). Park records 

are from the Páramo de Guaramacal in dwarf forest and on 

shrubs in areas dominated by bamboo; 2,900–3,000 m.

Brassia R. Br.

Brassia R. Br., in W. T. Aiton, Hort. Kew., 2nd ed., 5: 215. 1813.

Brachtia Rchb. f., Linnaea 22: 853. 1849, nom. cons.

Mesospinidium Rchb. f., Bot. Zeitung (Berlin) 10: 929. 1852.

Oncodia Lindl., Fol. Orchid. 4: Oncodia 1. 1853.

Ada Lindl., Fol. Orchid. 5: Ada 1. 1854.

Brassiopsis Szlach. & Górniak, Biodivers. Res. Conservation 1–2: 12. 2006.

Epiphytic herbs; caespitose or with shortly creeping, ascending rhizomes. Roots dense, thick. Pseudobulbs conspicuous, 

laterally compressed, apically 1- or 2-leaved, subtended by 1–4 

leaf-bearing sheaths. Leaves articulate; leaf blades conduplicate, 

usually elliptic or oblong-elliptic, apices acute or obtuse, coriaceous or chartaceous, epetiolate. Inflorescences 1 or 2, lateral, 

from the base of pseudobulbs and emerging among sheaths 

that subtend them or from the newly forming shoots, ± as long 

or longer than the leaves that subtend them, erect or arcuate, 

racemose with the flowers in the upper 1⁄3; peduncles remotely 

MAP 121. Brachionidium tuberculatum occurrence in Venezuela.
124 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

few-sheathed; rachises abbreviated; floral bracts glumaceous, 

stiff, distichous or unilateral, as long or longer than the pedicellate ovary, often covering the base of the flower. Flowers usually 

resupinate, secund on the inflorescence, cream, yellow, orangeyellow or greenish-yellow, often with a white labellum; perianth segments free, fleshy, ringent or campanulate. Sepals equal 

or dorsal sepal somewhat broader and lateral sepals slightly 

oblique, elliptic, ovate-elliptic or oblong-elliptic. Petals broader 

than the sepals, often triangular-elliptic. Labellum very fleshy, 

simple, concave, apical portion sigmoid or reflexed, basal portion calceolate, firmly adnate and confluent with the canaliculate ovary, disk provided with a pair of fleshy callus-like ridges 

below which there is a pad-like, pubescent or pilose callus at the 

entrance to the canaliculate nectary; column short, stout, parallel to the labellum, apically produced into a pair of short, thick 

wings, footless; stigmatic surface subreniform; rostellum bidentate; clinandrium smooth; anther smooth; pollinia 2, pyriform, 

stipes broad, subquadrate, viscidium suborbicular, relatively 

small. Ovary pedicellate, glabrous. Capsules 3-nerved. [Epidendroideae: Cymbidieae: Oncidiinae.]

A subtropical and tropical American genus of ~75 species 

found in North America (USA), Mexico, Central America, the 

West Indies, and South America (Colombia, Venezuela, Trinidad 

and Tobago, the Guianas, Ecuador, Peru, Bolivia, and Brazil). 

Fifteen species are found in Venezuela.

References. Chase and Whitten (2011); Pridgeon 

et al. (2009); Senghas (1999); Sweet (1974).

Brassia sulphurea (Rchb. f.) M. W. Chase, Phytotaxa 20: 28. 

2011. Brachtia sulphurea Rchb. f., Bonplandia (Hannover) 

2: 14. 1854; Foldats, Fl. Venez. 15(5): 57–59, fig. 786. 

1970; Ortega et al., BioLlania 5: 46. 1987.

Brachtia glumacea Rchb. f., Linnaea 22: 854. 1849; Dunsterville and Garay, 

Venez. Orchids Ill. 3: 44–45. 1965; Foldats, Fl. Venez. 15(5): 55–57, 

fig. 785. 1970; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 37. 

2000; Dorr et al., Contr. U.S. Natl. Herb. 40: 48. 2000 [2001]; non 

Brassia glumacea Lindl., Orchid Linden. 17. 1846. Oncodia glumacea

(Rchb. f.) Lindl., Fol. Orchid. 4: Oncodia 1. 1853. Brassiopsis glumacea (Rchb. f.) Szlach. & Górniak, Biodivers. Res. Conservation 1–2: 

12. 2006. Brassia glumaceoides M. W. Chase, Phytotaxa 20: 28. 2011, 

nom. nov.

Epiphytic herbs, 15–25 cm tall; pseudobulbs apically 1- or 

rarely 2-leaved, 2–3.5 cm long. Leaf blades 10–22 × 2 cm. Floral 

bracts ~2 cm long. Flowers fleshy and thick, bright yellow. Dorsal sepal 6–7.5 mm long.

Found in South America (Colombia and Venezuela). Venezuelan records are from the Cordillera de Mérida (Mérida and 

Trujillo) and the Cordillera de la Costa (Aragua; Map 122). 

Found in cloud forest and at forest margins along the Boconó–

Guaramacal road; 2,550–2,800 m.

Camaridium Lindl.

Camaridium Lindl., Bot. Reg. 10: t. 844. 1824. Maxillaria sect. Camaridium

(Lindl.) Christenson, Proc. 16th World Orchid Conf. 282. 2002.

Psittacoglossum La Llave & Lex., Nov. Veg. Descr. 2 (Orch. Opusc.): 29. 

1825.

Sepalosaccus Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19: 244. 1923.

Maxillaria sect. Cucullatae Christenson, Proc. 16th World Orchid Conf. 283. 

2002.

Pseudomaxillaria Hoehne, Arq. Bot. Estado São Paulo, ser. 2, 2: 71. 1947. 

Maxillaria sect. Pseudomaxillaria Christenson, Proc. 16th World Orchid Conf. 285. 2002.

Adamanthus Szlach., in Szlachetko and Śmiszek, Richardiana 7(1): 30. 2007.

Chaseopsis Szlach. & Sitko, in Szlachetko et al., Biodivers. Res. Conservation 

25: 25. 2012.

Chelyella Szlach. & Sitko, in Szlachetko et al., Biodivers. Res. Conservation 

25: 25. 2012.

Viracocha Szlach. & Sitko, in Szlachetko et al., Biodivers. Res. Conservation 

25: 36. 2012.

Epiphytic, rarely lithophytic or subterrestrial herbs; erect, 

creeping or pendulous; sympodial or pseudomonopodial; caespitose or rhizomatous. Vegetative architecture quite variable; in 

some species sympodia consist of a few short internodes, one of 

which is thickened into a ± conspicuous pseudobulb; in other 

species the sympodia consist of several extended erect, creeping 

or subpendulous internodes with at least 1 of the terminal internodes thickened into a pseudobulb; in other species pseudobulbs 

are absent, and the plant is essentially monopodial; in many 

species the juvenile plant is sympodial with aggregated pseudobulbs, and the mature plant produces new sympodia with long 

internodes; the long internodes generally covered with imbricate 

sheaths, with or without foliar blades. Pseudobulbs heteroblastic, 

1- or 2-foliate apically, subtended by several sheaths, of which 

the innermost 1 or 2 usually have leaf blades. Leaves conduplicate, articulate; leaf blades usually oblong or linear-oblong to 

MAP 122. Brassia sulphurea occurrence in Venezuela.
NUMBER 100 • 125

elliptic, symmetrical to irregularly oblique, sessile or sometimes 

pseudopetiolate. Inflorescences 1-flowered, solitary or fasciculate at the base of pseudobulbs or sheaths that cover the stems; 

peduncles relatively short, thin. Flowers usually resupinate, conspicuous and showy or inconspicuous and tiny, subcampanulate 

to spreading, usually short-lived, rarely flowers long-lasting, usually white or whitish-green, with or without nerves or spots of 

other colors, or rarely flowers of other colors entirely; perianth 

segments usually free, lacking conspicuous fibers, membranous, 

often minutely papillose with a sparkling texture, usually nectarless. Sepals subequal; lateral sepals often larger, oblique, adnate 

to the column foot, forming a mentum. Petals similar to sepals, 

but smaller. Labellum somewhat fleshier than other perianth 

parts, variable, entire or 3-lobed, wider than long or as wide as 

long, articulate to ± fused at the apex of the column foot; labellar disk lacking pseudopollen, wax or oil, provided with a single 

callus; column hemicylindrical or cylindrical, straight or curved 

with a short or almost absent foot; anther terminal, operculate, 

incumbent, often crested, glabrous; pollinia 4, in 2 unequal, 

superimposed pairs, stipe usually lunate, sometimes ligulate to 

seemingly absent; viscidium well-developed, broadly lunate or 

ovoid, rostellum transverse; stigma ventral, ovoid or transverse. 

Ovary smooth, round in cross section, pedicellate. Capsules ellipsoid to subspherical; valves separating apically, reflexing at 

dehiscence. [Epidendroideae: Cymbidieae: Maxillariinae.]

A neotropical genus of 80–85 species found in North America (USA), Mexico, Central America, the Greater Antilles, and 

South America (Colombia, Venezuela, Trinidad and Tobago, Ecuador, Peru, Bolivia, and Brazil). Camaridium has a center of diversity in Central America. Five or 6 species occur in Venezuela.

References. Blanco et al. (2007); Christenson 

(2002b); Pridgeon et al. (2009); Whitten et al. (2007).

Maxillaria s.l. is grossly polyphyletic (Blanco et al., 2007; 

Whitten et al., 2007) and Camaridium has been resurrected for 

members of the “core Maxillariinae” that have membranous, 

short-lived flowers that are basically white but often have red 

or purple nerves or spots. The flowers of Camaridium are subtended by conspicuous floral bracts, and the plants tend to have 

elongate stems and rhizomes. The pseudobulbs of Camaridium

are 1- or 2-foliate, whereas those of most species of Maxillaria

s. str. are 1-foliate.

We disagree with the taxonomy of Szlachetko et al. (2012), 

who split Camaridium into seven genera, and we have adopted a 

more conservative circumscription of the genus (see Blanco et al., 

2007; Whitten et al., 2007; Pridgeon et al., 2009).

Camaridium vestitum (Sw.) Lindl., Ann. Mag. Nat. Hist., ser. 3, 

1: 333. 1858. Epidendrum vestitum Sw., Prodr. 124. 1788. 

Camaridium purpureum Spreng., Syst. Veg. 3: 735. 1826, 

nom. illeg. Ornithidium vestitum (Sw.) Rchb. f., Ann. Bot. 

Syst. 6: 491. 1863. Ornithidium confertum Griseb., Fl. Br. W. 

I. 626. 1864. Maxillaria purpurea (Spreng.) Ames & Correll, 

Bot. Mus. Leafl. 11: 16. 1943; Dunsterville and Garay, Venez. 

Orchids Ill. 1: 234–235. 1959; Foldats, Fl. Venez. 15(4): 511–

514, fig. 751. 1970. Maxillaria conferta (Griseb.) C. Schweinf. 

ex León, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 

8: 395. 1946 [1947]; Romero and Carnevali, Orchids Venez., 

2nd ed., 2: 532. 2000; Carnevali and Ramírez-Morillo, in 

Berry et al., Fl. Venez. Guayana 7: 436. 2003; Fernández, 

Orquídeas Nat. Táchira 142. 2003. Pseudomaxillaria vestita

(Sw.) Brieger, Bot. Jahrb. Syst. 97: 553. 1977.

Scaphyglottis parviflora Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 58, t. 97. 1835 

[1836], non Camaridium parviflorum Fawc., in Urban, Symb. Antill. 

1: 472. 1910. Ornithidium parviflorum (Poepp. & Endl.) Rchb. f., 

Bonplandia (Hannover) 2: 19. 1854. Maxillaria parviflora (Poepp. & 

Endl.) Garay, Bot. Mus. Leafl. 21: 258. 1967; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 51. 2000 [2001]. Pseudomaxillaria parviflora (Poepp. 

& Endl.) Brieger, Bot. Jahrb. Syst. 97: 555. 1977. Camaridium micranthum M. A. Blanco, Lankesteriana 7: 520. 2007, nom. nov.

Epiphytic herbs, suberect or ascending to subpendulous; 

sympodia consisting of several internodes covered by scarious 

sheaths lacking leaf blades and terminating in a 1-foliate pseudobulb. Leaf blades elliptic, 6–19 × 0.6–1.5 cm, apices acute. 

Flowers white, campanulate. Perianth segments 4–6 mm long.

Found in North America (USA), Central America, the 

Greater Antilles, and South America (Colombia, Venezuela, Trinidad and Tobago, the Guianas, Ecuador, including the Galapagos Islands, Peru, Bolivia, Brazil, and Argentina). In Venezuela, 

known from the Andes (Portuguesa and Táchira), the Cordillera 

de la Costa (Aragua, Distrito Federal, Miranda, Monagas, and 

Yaracuy), and the Venezuelan Guayana (Amazonas, Bolívar, and 

Delta Amacuro; Map 123). Near La Divisoria de la Concepción; 

(1,300–)1,500–1,700 m.

A great deal of nomenclatural confusion surrounds this 

species. Venezuelan floras and checklists, with the exception of 

MAP 123. Camaridium vestitum occurrence in Venezuela.
126 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Knuth (1928: 267), generally have failed to appreciate either 

that Camaridium purpureum Spreng. is an illegitimate renaming 

of Epidendrum vestitum Lindl. or that Ornithidium confertum

Griseb. was proposed as a new name for E. vestitum Lindl. and 

that these names all share the same type and are therefore obligate synonyms.

Cleistes Rich. ex Lindl.

Cleistes Rich. ex Lindl., Gen. Sp. Orchid. Pl. 409. 1840.

Terrestrial or humicolous herbs; erect. Roots elongate, fibrous, fleshy or tuberous. Stems unbranched or rarely branching, terete or subcylindrical. Leaves convolute, not articulate, 

distichous to ± spiraled; leaf sheaths clasping; leaf blades 1 to 

several per stem or blades sometimes reduced and bract-like or 

totally absent, leaves or entire plant frequently glaucous, subfleshy to rarely subcoriaceous. Inflorescences terminal, racemose, few-flowered or 1-flowered and axillary at base of leaflike 

bracts. Flowers resupinate, short-lived, usually showy, greenish, 

yellow, white, purple, pink or red; floral bracts similar to leaf 

blades but smaller; perianth segments free, mostly campanulate, 

membranous to subfleshy. Sepals usually narrowly elliptic or 

narrowly oblong-elliptic, lateral sepals somewhat oblique, sometimes widely spreading. Petals similar to sepals, often wider and 

shorter or differently colored, parallel to column. Labellum free 

from column and parallel to it, basally with 2–4 pedunculate 

calli, entire or 3-lobed, flat or more frequently concave, inner 

surface generally with well-developed longitudinal calli or lamellae, frequently tuberculate; column relatively long, erect or forming an angle with respect to ovary, apically lobed or unlobed, 

usually tapering basally, footless; anther terminal, incumbent; 

clinandrium mostly 3-lobed or otherwise lobed or lacerate; pollinia 2, pollen soft and mealy, coherent in tetrads, without viscidium; rostellum transverse; stigma entire, ventral. Ovary relatively 

long, subterete, pedicellate, glabrous. Capsules ellipsoid, erect. 

[Vanilloideae: Pogonieae.]

A neotropical genus of ~30 species found in Central America (Costa Rica and Panama) and South America (Colombia, 

Venezuela, the Guianas, Ecuador, Peru, Brazil, Paraguay, and 

Argentina); Brazil is the center of species diversity. Ten species 

occur in Venezuela.

References. Pansarin and Barros (2008); Pansarin 

et al. (2008); Pridgeon et al. (2003).

Species of Cleistes are mostly sun loving and often found in 

boggy or marshy areas.

Authorship of the generic name has been attributed incorrectly to L. C. Richard. The confusion apparently stems from 

Lindley’s citation of “Cleistes. Rich. Annot. p. 9” in his protologue. However, the Richard (1817) publication that he cited 

does not contain a validating description, and none was supplied 

until Lindley adopted Richard’s generic name.

Molecular evidence (Pansarin et al., 2008) indicates that 

the genus Cleistes s.l. is paraphyletic and that the two (one?) 

North American species are more closely related to Isotria Raf. 

and Pogonia Juss. than they are to their Central American and 

South American congeners, which form a monophyletic group 

that includes the type of the genus. Pansarin and Barros (2008), 

citing molecular and morphological evidence, segregated the 

North American species as Cleistesiopsis Pansarin & F. Barros; 

the neotropical elements retain the name Cleistes.

Cleistes rosea Lindl., Gen. Sp. Orchid. Pl. 410. 1840; Dunsterville and Garay, Venez. Orchids Ill. 1: 82. 1959; Romero and 

Carnevali, Orchids Venez., 2nd ed., 1: 113. 2000; Carnevali 

and Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 7: 

279. 2003; Fernández, Orquídeas Nat. Táchira 45. 2003. 

Pogonia rosea (Lindl.) Rchb. f., Xenia Orchid. 2: 89. 1865; 

Foldats, Fl. Venez. 15(1): 137–139, fig. 46. 1969.

Terrestrial herbs, (30–)50–100 cm tall. Leaf blades 5–15 cm 

long. Flowers large and showy, ephemeral (lasting only 1 day), 

pink to purple, darker on the margins of the apical lobe. Dorsal 

sepal 6–8 cm long.

Found in Central America (Panama) and South America 

(Colombia, Venezuela, Trinidad and Tobago, the Guianas, Ecuador, Peru, Bolivia, and Brazil). In Venezuela, found in the Andes 

(Mérida, Táchira, and Trujillo), the Cordillera de la Costa (Aragua, Distrito Federal, Miranda, and Yaracuy), and the Venezuelan Guayana (Amazonas and Bolívar; Map 124). Park records 

are from the north slope of Guaramacal; 1,900–2,100 m.

This is the most widespread species of the genus. Although 

several of our references include Costa Rica in its distribution, 

Costa Rican material of Cleistes rosea has been segregated as 

C. costaricensis Christenson. Similarly, although two color forms 

MAP 124. Cleistes rosea occurrence in Venezuela.
NUMBER 100 • 127

of C. rosea once were recognized in Venezuela, C. rosea f. pallida

Carnevali & I. Ramírez has been described as a distinct species, 

C. abdita G. A. Romero & Carnevali, that is restricted to savannas in the Venezuelan Guayana. Cleistes abdita differs from our 

species in having white or pale pink (vs. pink, purple or deep 

purple) flowers and a narrower, denticulate (vs. entire) labellum.

Comparettia Poepp. & Endl.

Comparettia Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 42. 1835 [1836].

Diadenium Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 41. 1835 [1836].

Chaenanthe Lindl., Edwards’s Bot. Reg. 24: Misc. 38. 1838 (“Chænanthe”).

Scelochilus Klotzsch, Allg. Gartenzeitung. 9: 261. 1841.

Neokoehleria Schltr., Repert. Spec. Nov. Regni Veg. 10: 390. 1912.

Scelochiloides Dodson & M. W. Chase, Icon. Pl. Trop., ser. 2, 3: t. 293. 1989.

Stigmatorthos M. W. Chase & D. E. Benn., Lindleyana 8: 4. 1993.

Pfitzeria Senghas, J. Orchideenfreund 5(1): 20. 1998.

Scelochilopsis Dodson & M. W. Chase, Orquideología 21: 61. 1998.

Epiphytic herbs; erect or pendulous; caespitose; sometimes 

partially psygmoid (i.e., with laterally compressed leaves that 

are arranged fanlike); sympodial. Rhizomes short or absent; 

pseudobulbs inconspicuous to almost absent, usually cylindrical to suborbicular, apically 1-leaved, enclosed by 1 or 2(3 or 

more) overlapping sheaths, of which the innermost 1 or 2 have 

leaf blades (sometimes absent). Leaves conduplicate or less 

commonly laterally compressed, articulate, usually oblong or 

oblong-elliptic, acute or obtuse, often tinged with red, sessile or 

short-petiolate. Inflorescences arising from the base of pseudobulbs, erect, arching or pendulous, shorter to longer than the 

leaves, usually in lax, few-flowered racemes, sometimes a lateral 

branch arises from the base of the rachis or from the older internodes of the peduncle; peduncles glabrous, with remote tubular 

sheaths; floral bracts inconspicuous, tubular to concave. Flowers resupinate or not, small or medium, long-pedicellate, usually 

relatively short-lived, yellow, yellow with reddish nerves, red, 

orange or pinkish with darker purple nerves; perianth segments 

parallel to column at base, apical 1⁄2 straight or spreading, membranous to subfleshy. Dorsal sepal free, usually concave; lateral 

sepals free apically or not, connate basally and forming a spur 

of various shapes, apically 1- or 2-lobed. Petals usually broader 

than the sepals, free. Labellum more conspicuous than other 

perianth segments, hastate basally and enclosed in the sepaline 

spur, lamina simple, variable in shape, usually 2-lobed or apically emarginate; disk with callus consisting of 2 parallel longitudinal keels; column erect, straight; anther terminal with 

a basal horn or pair of horns that secrete nectar into a cavity 

formed by fused lateral sepals, not auriculate or with very small 

wings, almost footless; pollinarium with a flat, spathulate stipe 

longer than the pollinia; caudicles very small; pollinia 2, yellow, 

orbicular, pyriform to flattened and disciform; viscidium circular, 

entire, ventral; rostellum beak-shaped; stigmatic surface immediately below the rostellum, circular or in the form of a small slit. 

Ovary glabrous. Capsules ellipsoid, triangular. [Epidendroideae: 

Cymbidieae: Oncidiinae.]

A neotropical genus of ~60 species found in Mexico, Central America, the Greater Antilles, and South America (Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, and Brazil); 

most diverse in the western Andes. Six species occur in Venezuela. Several species, especially species of Comparettia s. str., also 

are cultivated in Venezuela.

References. Bock (1986); Bockemühl and Senghas 

(1985); Neubig et al. (2012); Pridgeon et al. (2009); Senghas 

(1987, 1995d, 1995e, 2001); Tuskes and Tuskes (1999).

Comparettia species are usually sun-loving twig epiphytes, 

and many of them are quite inconspicuous, infrequently collected, and, as a consequence, poorly known.

We have adopted an expanded circumscription of this genus 

based on molecular evidence (Pridgeon et al., 2009; Neubig et 

al., 2012). Comparettia s.l. now includes all species with sepalar 

nectar spur(s) furnished by a horn or pair of horns on the column 

base that secrete nectar. These species previously were scattered 

among the genera listed in synonymy.

KEY TO THE SPECIES OF COMPARETTIA

1a. Inflorescences usually longer than the leaves; flowers lilac or reddish; pedicels longer than dorsal sepal; labellum conspicuously larger than other perianth segments, widely diverging, apical lobe wider than long; nectary >1 cm long

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. falcata

1b. Inflorescences usually shorter or equal in length to the leaves; flowers yellow with reddish nerves; pedicels shorter than 

dorsal sepal; labellum ± the same length as petals and sepals and enclosed in a tubular flower, apical lobe longer than 

wide; nectary 0.3–0.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ottonis

Comparettia falcata Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 42, t. 

73. 1835 [1836]; Dunsterville and Garay, Venez. Orchids 

Ill. 1: 86–87. 1959; Foldats, Fl. Venez. 15(5): 89–92, fig. 

795. 1970; Romero and Carnevali, Orchids Venez., 2nd ed., 

1: 123. 2000; Carnevali and Ramírez-Morillo, in Berry et 

al., Fl. Venez. Guayana 7: 284. 2003; Fernández, Orquídeas 

Nat. Táchira 47. 2003.

The nominate variety is the only one found in Guaramacal 

and in Venezuela.

Comparettia falcata var. falcata

Twig epiphytes. Leaf blades 5–20 cm long. Inflorescences 

(5–)10–20(–30) cm long. Flowers lilac or reddish. Labellum 1–2 

cm wide.
128 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Found in Mexico, Central America, the Greater Antilles, and tropical South America (Colombia, Venezuela, French 

Guiana, Ecuador, Peru, Bolivia, and Brazil). Widely distributed 

in Venezuela (Amazonas, Aragua, Bolívar, Carabobo, Distrito 

Federal, Falcón, Lara, Mérida, Miranda, Portuguesa, Táchira, 

Trujillo, and Yaracuy; Map 125). Park records are from near 

the Laguna de Aguas Negras and in the Qda. Chandá; (1,350–)

1,800–1,850 m.

A collection of this species made near the lower elevational 

boundary of the park was described as having white flowers 

(Angulo & Infante 2, PORT), but this is probably an observational error made in the field. Flower color in this species 

suggests hummingbird pollination, which is what has been observed in plants growing in Puerto Rico (Rodríguez-Robles et 

al., 1992).

Brazilian material of this species is segregated as Comparettia falcata var. paulensis (Cogn.) I. Bock.

Comparettia ottonis (Klotzsch) M. W. Chase & N. H. Williams, 

in Chase et al., Lindleyana 21: 29. 2008. Scelochilus ottonis Klotzsch, Allg. Gartenzeitung. 9: 261. 1841; Foldats, 

Fl. Venez. 15(5): 79–81, fig. 792. 1970; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 964. 2000; Dorr et al., 

Contr. U.S. Natl. Herb. 40: 54. 2000 [2001]; Carnevali and 

Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 7: 561. 

2003; Fernández, Orquídeas Nat. Táchira 215. 2003.

Rodriguezia stenochila Lindl., Orchid. Linden. 23. 1846. Scelochilus stenochilus (Lindl.) Rchb. f., Ann. Bot. Syst. 6: 689. 1861 [1863]; Foldats, Fl. 

Venez. 15(5): 82. 1970. Comparettia stenochila (Lindl.) M. W. Chase 

& N. H. Williams, in Chase et al., Lindleyana 21: 30. 2008.

Twig epiphytes. Leaf blades 8–14 cm long. Inflorescences ± 

the same length as the leaves. Flowers golden-yellow, red striped 

or brown. Labellum golden-yellow with red-purple areas, 1.2–

1.7 cm long.

Restricted to Venezuela, although perhaps also present in 

the Brazilian portion of the Sierra de la Neblina. In Venezuela, 

collected in the Andes (Mérida, Táchira, and Trujillo), the Cordillera de la Costa (Aragua, Carabobo, Distrito Federal, Miranda, and Yaracuy), and the Venezuelan Guayana (Sierra de la 

Neblina, Amazonas state; Map 126). Locally common in montane forest on the north slope of the park and between Boconó 

and Batatal; 1,950–2,600 m.

Rodriguezia stenochila Lindl., described from nearby 

Mérida state and only known from the type, was thought to 

differ from Comparettia ottonis by a narrower, apically hastate 

labellum. Examination of an image of the type of R. stenochila

suggests that this species was most likely based on an artifact 

created by the poor preservation of its flowers, which is not surprising as flowers of Comparettia are membranous and difficult 

to reconstruct after being pressed and dried.

Corymborkis Thouars

Corymborkis Thouars, Nouv. Bull. Sci. Soc. Philom. Paris, sér. 2, 1: 318. 

1809.

Terrestrial herbs; erect; caespitose; sympodial. Roots few, 

fibrous, fascicled. Rhizomes short, subterranean. Stem cane-like, 

somewhat woody basally, usually unbranched, to several meters 

long, covered by leaf sheaths. Leaves distichous or sometimes 

spiral, few to many, plicate, with several prominent nerves, not 

MAP 125. Comparettia falcata var. falcata occurrence in Venezuela. MAP 126. Comparettia ottonis occurrence in Venezuela.
NUMBER 100 • 129

articulate; leaf sheaths tubular and almost completely covering 

the stem, often brown furfuraceous basally; leaf blades elliptic, 

acuminate. Inflorescences lateral or (more rarely) subterminal, 

erect or patent, much shorter than the subtending leaves, solitary in each leaf axil (the plant often with several inflorescences 

simultaneously), racemes or more commonly panicles that are 

1- or 2-pinnate with 3–10-flowered branches; peduncles terete 

or flattened, ± naked; rachises relatively short with subfoliaceous 

sheaths. Flowers distichous, resupinate or not; pedicels relatively 

short, sometimes ± showy and fragrant, campanulate; perianth 

segments subequal, linear-elliptic to linear-obovate, connivent or 

connate basally, sometimes clawed. Labellum with 2 conspicuous keels that completely encircle the column, leaving only the 

labellar apex free, which is somewhat reflexed, without a callus; 

column erect, nearly as long as the labellum; anther terminal, 

erect, pollinia 2, thin, sectile, bipartite, attached to a linear-ovate 

viscidium via an elongated hamulus (i.e., pollinium stalk derived 

from the apex of the rostellum), flat or semiterete; rostellum 

prominent, bifid when viscidium removed. Ovary cylindrical, 

glabrous. Capsules fusiform, 6-ribbed, crowned by the persistent perianth and the remnants of the column. [Epidendroideae: 

Tropidieae.]

A pantropical genus of 5 species, 3 of which occur in the 

neotropics. The American species are found in Mexico, Central 

America, the West Indies, and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Paraguay, and Argentina). 

Two species, one of them endemic, are known from Venezuela.

References. Pridgeon et al. (2005); Rasmussen 

(1977).

Species of Corymborkis are shade loving. 

The leaves on specimens of Corymborkis that have been 

prepared for the herbarium are typically dark reddish brown 

when dry.

Corymborkis flava (Sw.) Kuntze, Revis. Gen. Pl. 2: 658. 1891 

(“Corymborchis”); Foldats, Fl. Venez. 15(1): 237–239, fig. 

88. 1969; Dunsterville and Garay, Venez. Orchids Ill. 6: 94–

96. 1976; Romero and Carnevali, Orchids Venez., 2nd ed., 

1: 129. 2000 (“Corymborchis”). Serapias flava Sw., Prodr. 

119. 1788.

Terrestrial herbs, erect, 1–3 m tall. Leaf blades 20–30 × 5–7 

cm. Flowers campanulate, yellow, relatively attractive. Sepals 

1.4–1.7 cm long.

Found in Central America, the West Indies, and South 

America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, 

Paraguay, and Argentina). In Venezuela, known from the Andes 

(Lara and Trujillo) and the Cordillera de la Costa (Aragua, Distrito Federal, and Miranda; Map 127). In the park, found in 

Qda. Honda; ~1,900 m.

Corymborkis galipanensis (Rchb. f.) Foldats, which is 

endemic to the Cordillera de la Costa, can be distinguished 

from C. flava by its much longer (30 vs. 15 mm long) perianth 

segments.

Cranichis Sw.

Cranichis Sw., Prodr. 8, 120. 1788.

Terrestrial, humicolous or rarely subepiphytic herbs; caespitose. Roots fascicled, fleshy, villous. Leaves usually basal, often in 

a rosette, rarely cauline and gradually intergrading into peduncular sheaths, usually present at anthesis, often with white spots, 

herbaceous to subfleshy, sessile or petiolate. Inflorescences terminal, racemose or pseudospicate, usually lax; peduncles terete, with 

few tubular sheaths. Flowers not resupinate, small; perianth segments usually green or white, membranous, labellum often with 

conspicuous, prominent, reticulate greenish or purplish veins. Sepals free. Petals free, much narrower than sepals. Labellum fleshier 

than other perianth segments, usually concave or cochleariform, 

sometimes lobed; column fleshy, short; clinandrium cyathiform, 

acute, erect; stigma relatively large, terminal; anther dorsal, erect; 

pollinia 4, ovoid-clavate, with a small, brittle hamulus; rostellum 

apical, viscidium small, globose. Ovary long-pedicellate, pubescent. Capsules erect. [Orchidoideae: Cranichideae: Cranichidinae.]

A subtropical and tropical American genus of ~60 species 

found in North America (USA), Mexico, Central America, the 

West Indies, and South America (Colombia, Venezuela, Suriname, French Guiana, Ecuador, Peru, Bolivia, Brazil, and Argentina). Approximately a dozen species occur in Venezuela, the 

majority in the Andes.

References. Álvarez-Molina and Cameron (2009); 

Pridgeon et al. (2003); Salazar Chávez et al. (2009).

Cranichis appears to be monophyletic (Álvarez-Molina and 

Cameron, 2009; Salazar Chávez et al., 2009) and is part of a 

larger, well-supported clade comprising the Cranichidinae s. str.

MAP 127. Corymborkis flava occurrence in Venezuela.
130 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

KEY TO THE SPECIES OF CRANICHIS

1a. Leaves long-petiolate, petioles longer than or ± equal to leaf blades, >5 cm long; labellum suborbicular or obovate, with 

thick, elevated, green or purple longitudinal ribs; petal margins ciliate . . . . . . . . . . . . . . . . . . . . . . . . . C. antioquiensis

1b. Leaves short-petiolate, petioles shorter than leaf blades, ≤5 cm long; labellum ovate, without thick, longitudinal ribs but 

with a large, 2-lobed, ± verruculose callus; petal margins not ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. diphylla

Cranichis antioquiensis Schltr., Repert. Spec. Nov. Regni Veg. 

Beih. 7: 57. 1920; Romero and Carnevali, Orchids Venez., 

2nd ed., 1: 130. 2000; Fernández, Orquídeas Nat. Táchira 

47. 2003; Morillo, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 1: 103, fig. 6. 2011.

Cranichis ciliata auct., non (Kunth) Kunth; Dunsterville and Garay, Venez. 

Orchids Ill. 3: 66–67. 1965, pro parte; Foldats, Fl. Venez. 15(1): 410–

411, fig. 153. 1969, pro parte; Dorr et al., Contr. U.S. Natl. Herb. 40: 

49. 2000 [2001].

Terrestrial herbs, to 45 cm tall. Leaf blades 3–15 × 2–7(–8) 

cm. Inflorescences to 20 cm tall. Flowers green or white tinged 

with dark green or brown. Dorsal sepal 4.5–6 mm long. Labellum white with raised, deep green or brown ribs.

Found in South America (Colombia, Venezuela, and Ecuador). In Venezuela, restricted to the Andes (Lara, Mérida, 

Táchira, and Trujillo; Map 128). In the park, found near the Laguna de Aguas Negras and along the Boconó–Guaramacal road; 

1,850–2,150 m.

This species has been confused with Cranichis ciliata

(Kunth) Kunth (described from the Silla de Caracas in the Distrito Federal), but it has larger flowers, a narrower labellum, and 

inflorescences with more flowers. It may be, however, that these 

two species are not distinct, but that question requires further 

study.

Cranichis diphylla Sw., Prodr. 120. 1788; Foldats, Fl. Venez. 

15(1): 411–413, fig. 154. 1969; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 131. 2000; Morillo, in Morillo et 

al., Monocot. Páramos Venez. 1: 104–105, fig. 7. 2011.

Cranichis monophylla Lindl., Orchid. Linden. 27. 1846; Dunsterville and 

Garay, Venez. Orchids Ill. 4: 58–59. 1966; Carnevali and RamírezMorillo, in Berry et al., Fl. Venez. Guayana 7: 257, fig. 250. 2003.

Cranichis tenuis auct., non Rchb. f.; Dunsterville, Lindleyana 1: 234–235. 

1986; Ortega et al., BioLlania 5: 46. 1987; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 133. 2000; Dorr et al., Contr. U.S. Natl. Herb. 

40: 49. 2000 [2001]; Fernández, Orquídeas Nat. Táchira 48. 2003.

Habenaria parviflora auct., non Lindl.; Dorr et al., Contr. U.S. Natl. Herb. 

40: 49. 2000 [2001] (as to Stergios & Palacios 18703).

Terrestrial herbs, 15–25(–30) cm tall. Leaf blades 3–6 × 

2–5(–8) cm, often variegated. Inflorescences to 20–27 cm tall. 

Flowers white with green veins. Dorsal sepal 1.5–3.5 mm long. 

Labellum white with some bright green spots.

Found in Mexico, Central America, the Greater Antilles, 

and South America (Colombia, Venezuela, Suriname, Ecuador, 

and Peru). Venezuelan records are from the Andes (Mérida, 

Táchira, and Trujillo), the Cordillera de la Costa (Aragua, 

Distrito Federal, and Miranda), and the Venezuelan Guayana 

(Bolívar; Map 129). In the park, found in Qda. Segovia along 

the Boconó–Guaramacal road; 2,000–2,400 m.

MAP 128. Cranichis antioquiensis occurrence in Venezuela. MAP 129. Cranichis diphylla occurrence in Venezuela.
NUMBER 100 • 131

Crocodeilanthe Rchb. f. & Warsz.

Crocodeilanthe Rchb. f. & Warsz., Xenia Orchid. 1: 10. [1 Apr] 1854; ibid., 

Bonplandia (Hannover) 2: 113. [1 Mai] 1854. Pleurothallis subgen. 

Crocodeilanthe (Rchb. f. & Warsz.) Luer, Monogr. Syst. Bot. Missouri 

Bot. Gard. 20: 34. 1986.

Stelis auct., non Sw.; Pridgeon and M. W. Chase, Lindleyana 16: 235–271. 

2001, pro parte, excluding type.

Epiphytic or lithophytic herbs; erect, large and robust (in 

comparison to other Pleurothallidinae). Rhizomes often abbreviated, less commonly creeping. Stems robust (in comparison to 

other Pleurothallidinae), elongate, conspicuously longer than the 

leaves to ± similar in length, terete, often partially covered by 1 to 

several appressed, imbricate tubular sheaths, basally covered in 

several coarse, papery sheaths, these often covering >1 stem at a 

time, stems in several species prolific (i.e., 1 or more stems originating from the apex of another). Leaves rarely erect with respect 

to the stem; leaf blades usually elliptic or obovate, bases petiolate 

or sessile, apices minutely 3-toothed, coriaceous, fleshy or fleshycoriaceous. Inflorescences terminal or emerging below the stem 

apex, solitary or often fasciculate, erect, subtended by a conspicuous foliaceous spathe associated with an annulus and emerging 

just above the annulus and below the leaf articulation, racemose, 

usually many-flowered, flowers simultaneous or rarely ± successive, shorter than subtending leaves to several times as long; floral 

bracts usually reduced or rarely very conspicuous, membranous; 

pedicels thick and shorter than or equal to the ovary. Flowers 

commonly resupinate, small or medium in size, erect, patent or 

nutant, variously colored but typically translucent, white, greenish or yellowish, with or without purple or brown veins, rarely 

entirely reddish-brown, membranous. Sepals almost free, fused 

basally or lateral for variable lengths (rarely completely fused) 

in a flat synsepal, bases concave, apices convex, never caudate, 

smooth, pubescent or verruculose internally, smooth externally; 

dorsal sepal often a different width than synsepal, apices acute 

to rounded. Petals conspicuously smaller and less fleshy than 

sepals, 1- or 3-nerved, variable in shape but usually oblong to 

elliptic or obovate, at least the basal 1⁄2 parallel to the column, 

margins smooth. Labellum supported by or contained within the 

concavity of the synsepal, articulate with the foot of the hingeshaped column (the basal part of the labellar blade is adaxially 

concave, and the concavity is freely articulated on the bulbous 

columnar foot), sometimes the hinged articulation is vestigial, 

and the labellum is merely articulate with the column base via 

an area of membranous tissue, without lobes or retrorse teeth 

basally, with a linear, transverse callus that separates the convex area that is articulated with the base of the labellar blade, 

the blade from almost simple to more commonly 3- or 5-lobed, 

the lateral lobes relatively small, erect or extended, apical lobe 

smooth, verruculose or pubescent, margins smooth, less commonly erose, disk usually with 2 calli, globose in the middle or on 

the side lobes; column relatively short, cylindrical, nearly straight 

or ± curved, with a well-developed, bulbous foot, smooth or verruculose; clinandrium usually entire; anther apical or subapical, 

operculate; stigma and rostellum ventral; pollinia 2. Ovary articulate with pedicel, terete or subangular. Capsules often ellipsoid 

or obclavate. [Epidendroideae: Epidendreae: Pleurothallidinae.]

A neotropical genus of 75–80 species found in Mexico, Central America, the West Indies, and South America (Colombia, 

Venezuela, Ecuador, Peru, and Brazil). The majority of species 

are found in the northern Andes at higher elevations (above 

1,500 m); only 1 species occurs in the West Indies. Approximately 10–12 species are reported from Venezuela.

References. Karremans et al. (2013); Luer (1986d, 

1998); Pridgeon and Chase (2001).

In Venezuela, Crocodeilanthe traditionally has been treated 

as part of Pleurothallis (see e.g., Foldats, 1970a; Dunsterville 

and Garay, 1979c; Romero-González and Carnevali FernándezConcha, 2000). Molecular phylogenetic studies, however, have 

provided a basis for reclassifying the Pleurothallidinae and contributed to a dismemberment of Pleurothallis s.l. (Pridgeon and 

Chase, 2001; Pridgeon et al., 2005). One result of these studies is the realization that Crocodeilanthe belongs to a clade that 

includes Stelis and several other groups previously included in 

Pleurothallis as subgenera or sections (see, e.g., Pridgeon and 

Chase, 2001; Pridgeon et al., 2005; Karremans et al., 2013). 

Pridgeon and Chase (2001) decided to treat this entire clade 

within a broadly circumscribed Stelis. Luer (2004, 2006, 2007, 

2011) rejected this broad circumscription of Stelis and instead 

recognized Stelis in a narrow sense, resurrecting older generic 

concepts such as Crocodeilanthe and creating a number of new 

genera to accommodate species that were excluded from Stelis

s. str. More recently, Karremans et al. (2013) identified a highly 

supported clade that includes both Stelis and Crocodeilanthe but 

one that also had four major subclades, including a subclade that 

corresponds to Stelis s. str. and another to Crocodeilanthe.

We have chosen to accept Crocodeilanthe because there is 

molecular support for a monophyletic lineage, and the genus 

is also well circumscribed morphologically. Species of Crocodeilanthe are relatively large plants with stems usually exceeding the leaves in length; sheaths enclosing the bases of stems; 

inflorescences multiflowered, erect, racemose and arising from 

conspicuous spathes; flowers resupinate, small, and opening simultaneously; sepals similar to each other and never caudate; 

petals 1- or 3-veined; labellum short and simple with 2 small 

lateral lobes attached to the base of the column; anther apical 

or subapical; and pollinia “bubble-like” (see Karremans et al., 

2013: fig. 8).

KEY TO THE SPECIES OF CROCODEILANTHE

1a. Rhizomes creeping; stems in fascicles of 2 or 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. stergiosii

1b. Rhizomes short, stems clustered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
132 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

2a. Leaves <1⁄2 the length of stems; inflorescences conspicuously shorter than leaves; flowers translucent, cream, green or 

white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Internodes of inflorescence rachis as long or almost as long as floral bracts; floral bracts ± inflated and as long as 

pedicels; sepals fused in the lower 1⁄2 of their length, synsepal geniculate; apices of petals and labellum rounded 

to subacute; labellum no more than 2× as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sp. A

 3b. Internodes of inflorescence rachis conspicuously longer than floral bracts; floral bracts appressed to the rachis 

and shorter than pedicels; sepals fused only in the lower 1

⁄5 of their length or nearly free, synsepal not geniculate; 

apices of petals and labellum truncate; labellum 2.5–3× longer than wide . . . . . . . . . . . . . . . . . . . . . C. gelida

2b. Leaves at least 3⁄4 the length of stems and often longer than stems; inflorescences ± equal in length to leaves or conspicuously longer than leaves; flowers usually translucent with purple or brown veins, rarely completely reddishbrown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Flowers (in living material) erect; peduncles very short, almost completely hidden by spathe . . . . . . C. moritzii

 4b. Flowers (in living material) nutant or patent; peduncles short or long, but always exceeding spathe in length

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 5a. Inflorescences slightly shorter to slightly longer than leaves, usually 3 or more present simultaneously; flowers (in living material) nutant; dorsal sepal 4–5 mm long, slightly wider than synsepal, ovate to elliptic ovate, 

3-ribbed, apex acute to broadly obtuse; petals spathulate or oblong-spathulate, 1-nerved; labellum elliptic in 

general outline, 3-lobed, with 2 subglobular calli, 1 on each side of the lateral lobe . . . . . . . . . . C. galeata

 5b. Inflorescences usually longer than leaves, 1 or 2 present simultaneously; flowers (in living material) patent; 

dorsal sepal 8–10 mm long, much wider than the synsepal, broadly ovate, 5-ribbed, apex rounded; petals 

broadly obovate, obovate or oblong, 3-nerved; labellum suborbicular or subquadrate-suborbicular in general outline, simple, ecallose or almost ecallose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. elegans

Crocodeilanthe elegans (Kunth) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95: 256. 2004; Morillo, in Morillo et al., 

Bot. Ecol. Monocot. Páramos Venez. 1: 106–107, fig. 8. 

2011. Dendrobium elegans Kunth, in H. B. K., Nov. Gen. 

Sp. [fol.] 1: 287. 1815 [1816]; ibid. [qu.] 1: 358. 1815 

[1816]. Pleurothallis elegans (Kunth) Lindl., Edwards’s Bot. 

Reg. 28: Misc. 70. 1842; Dunsterville and Garay, Venez. 

Orchids Ill. 4: 214–215. 1966; Foldats, Fl. Venez. 15(2): 

262–264, fig. 278. 1970; Romero and Carnevali, Orchids 

Venez., 2nd ed., 3: 778. 2000; Fernández, Orquídeas Nat. 

Táchira 180. 2003. Stelis elegans (Kunth) Pridgeon & M. W. 

Chase, Lindleyana 16: 262. 2001, non Luer & R. Vásquez, 

1981. Stelis picta Pridgeon & M. W. Chase, Lindleyana 17: 

100. 2002, nom. nov.

Epiphytic herbs, 15–30 cm tall (including inflorescences). 

Leaf blades narrowly elliptic, elliptic or oblong-elliptic, 9–13 cm 

long (including pseudopetioles), 1.6–5.5 cm wide. Inflorescences 

14–17 long, 12–26-flowered. Flowers pale brown with dark purple nerves. Dorsal sepal 8–10 mm long.

Found in South America (Colombia, Venezuela, Ecuador, and Peru). In Venezuela, known from the Andes (Mérida, 

Táchira, and Trujillo), the Cordillera de la Costa (Aragua and 

Distrito Federal), and the Venezuelan Guayana (Bolívar; Map 

130). In the park, known from the Páramo de Guaramacal near 

the television antennas; 2,900–3,100 m.

Crocodeilanthe galeata (Lindl.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95: 256. 2004; Morillo, in Morillo et al., 

Bot. Ecol. Monocot. Páramos Venez. 1: 109–110, fig. 10. 

2011. Pleurothallis galeata Lindl., Ann. Mag. Nat. Hist. 

15: 107. 1845; Dunsterville and Garay, Venez. Orchids Ill. 

4: 216–217. 1966; Foldats, Fl. Venez. 15(2): 282–283, fig. 

285. 1970; Ortega et al., BioLlania 5: 49. 1987; Dorr et al., 

Contr. U.S. Natl. Herb. 40: 53. 2000 [2001]; Romero and 

Carnevali, Orchids Venez., 2nd ed., 3: 788. 2000; Carnevali 

and Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 7: 

515. 2003; Fernández, Orquídeas Nat. Táchira 182. 2003. 

Stelis galeata (Lindl.) Pridgeon & M. W. Chase, Lindleyana 

16: 263. 2001.

MAP 130. Crocodeilanthe elegans occurrence in Venezuela.
NUMBER 100 • 133

Epiphytic or subterrestrial herbs, 20–30 cm tall (including 

inflorescences). Leaf blades elliptic, 9–15 cm long (including 

pseudopetioles), 2.5–3.5 cm wide. Inflorescences 10–15 cm long, 

30-flowered or more. Flowers cream-colored, pink or opaquetranslucent yellow to salmon, outer nerves a darker hue. Dorsal 

sepal 4.5–6 mm long.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela the distribution is disjunct, with records 

from the Andes (Apure, Lara, Mérida, Táchira, and Trujillo) and 

the Venezuelan Guayana (Amazonas and Bolívar; Map 131). In 

the park, collected in montane forest in various localities on both 

slopes of Guaramacal; 1,800–2,200(–2,600) m.

Crocodeilanthe gelida (Lindl.) Carnevali & I. Ramírez, comb. 

nov. Pleurothallis gelida Lindl., Edwards’s Bot. Reg. 27: 

Misc. 91. 1841; Foldats, Fl. Venez. 15(2): 283–285, fig. 

286. 1970; Ortega et al., BioLlania 5: 49. 1987; Romero 

and Carnevali, Orchids Venez., 2nd ed., 3: 790. 2000; Dorr 

et al., Contr. U.S. Natl. Herb. 40: 53. 2000 [2001]; Fernández, Orquídeas Nat. Táchira 182. 2003; Morillo, in Morillo 

et al., Bot. Ecol. Monocot. Páramos Venez. 1: 249–250, fig. 

94. 2011. Stelis gelida (Lindl.) Pridgeon & M. W. Chase, 

Lindleyana 16: 263. 2001. Specklinia gelida (Lindl.) Luer, 

Monogr. Syst. Bot. Missouri Bot. Gard. 95: 260. 2004. Niphantha gelida (Lindl.) Luer, Monogr. Syst. Bot. Missouri 

Bot. Gard. 120: 154. 2010.

Epiphytic herbs, 25–40 cm tall (including inflorescences). 

Leaf blades narrowly elliptic to oblanceolate, 8–12 cm long (including pseudopetioles), 3.3–3.8 cm wide. Inflorescences 6–9 cm 

long, 10–15-flowered. Flowers yellowish-green. Dorsal sepal 

5–8 mm long.

Found in North America (USA), Mexico, Central America, 

the Greater Antilles, and South America (Colombia, Venezuela, 

Ecuador, Peru, Bolivia, and Brazil). In Venezuela, collected in 

the Andes (Táchira and Trujillo) and the Cordillera de la Costa 

(Yaracuy; Map 132). In the park, known from the north slope of 

Guaramacal; 2,800 m.

The correct generic placement of this species is problematic, 

as its extensive synonymy suggests. We place it in Crocodeilanthe

on the basis of its morphology, although it is resolved in its own 

clade separate from Crocodeilanthe and Stelis s. str. in the molecular study published by Karremans et al. (2013; Clade E, but 

mislabeled D in fig. 5). Morphologically, C. galeata is very similar to their definition of Crocodeilanthe and Stelis s. str., except 

that this species has incumbent (vs. apical) anthers and divergent 

“whale tail” (vs. “bubble-like”) pollinaria (see Karremans et al., 

2013: fig. 8). Luer (2010) made this species the type of Niphantha Luer, which also includes a second species from Ecuador that 

has not been included yet in molecular analyses.

Crocodeilanthe moritzii (Rchb. f.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95: 256. 2004. Pleurothallis moritzii Rchb. 

f., Linnaea 22: 824. 1849; Dorr et al., Contr. U.S. Natl. 

Herb. 40: 53. 2000 [2001]; Carnevali and Ramírez-Morillo, 

in Berry et al., Fl. Venez. Guayana 7: 518–519, fig. 459. 

2003. Stelis moritzii (Rchb. f.) Pridgeon & M. W. Chase, 

Lindleyana 16: 264. 2001.

Pleurothallis elegans auct., non (Kunth) Lindl.; Foldats, Fl. Venez. 15(2): 

262–264, fig. 264. 1970.

Epiphytic, lithophytic or rarely subterrestrial herbs, 

15–30 cm tall (including inflorescences). Leaf blades elliptic 

to oblong-elliptic, 7–15 cm long (including pseudopetioles), 

MAP 131. Crocodeilanthe galeata occurrence in Venezuela. MAP 132. Crocodeilanthe gelida occurrence in Venezuela.
134 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

1.6–5.5 cm wide. Inflorescences 2–7 in a fascicle, 8–18 cm 

long, many-flowered. Flowers cream or yellowish. Dorsal sepal 

5–8 mm long.

Found in South America (Venezuela, Guyana, Ecuador, and 

Peru). In Venezuela, this species is reported from Amazonas, 

Apure, Aragua, Bolívar, Distrito Federal, Táchira, and Trujillo 

(Map 133). The sole park record is from subpáramo and dwarf 

forest below the Páramo de Guaramacal; 2,800 m.

♦ Crocodeilanthe stergiosii (Carnevali & I. Ramírez) Carnevali 

& I. Ramírez, in Hokche et al., Nuevo Cat. Fl. Vasc. Venez. 

578. 2008. Pleurothallis stergiosii Carnevali & I. Ramírez, 

Harvard Pap. Bot. 3: 247, figs. 5, 6. 1988; Ortega et al., 

BioLlania 5: 49. 1987; Dorr et al., Contr. U.S. Natl. Herb. 

40: 53. 2000 [2001].

Pleurothallis velaticaulis auct., non Rchb. f.; Foldats, Fl. Venez. 15(2): 440–

443, fig. 355. 1970, pro parte; Ortega et al., BioLlania 5: 49. 1987.

Epiphytic herbs, to 40 cm tall (including inflorescences); 

creeping. Leaf blades elliptic or oblong-elliptic, leaves 8.1–11.5 

cm long (including pseudopetioles), 1.5–3 cm wide. Inflorescences 2 or 3 in a fascicle, 15–24 cm long, much longer than 

the subtending leaves, 40–55-flowered. Flowers green or yellow. 

Dorsal sepal 4.5–5 mm long.

Endemic to Venezuela (Trujillo; Map 134). In the park, 

found on the north slope of Guaramacal; 1,800–2,700 m.

Crocodeilanthe sp. A

Epiphytic herbs, 28–35 cm tall (including inflorescences). 

Leaf blades lanceolate or narrowly elliptic, 13–15 cm long (including pseudopetioles), 2–2.8 cm wide. Inflorescences 5–7 cm 

long, 13–20-flowered. Flowers very small, greenish-white. Dorsal sepal 3.2–3.5 mm long.

Known only from the Andes of Venezuela (Trujillo; Map 

135). In the park, found on the north slope of Guaramacal near the Laguna de Aguas Negras and in Qda. Salvaje; 

1,850–2,100 m.

This appears to be an undescribed species currently known 

only from material collected in Guaramacal National Park.

MAP 135. Crocodeilanthe sp. A occurrence in Venezuela.

MAP 133. Crocodeilanthe moritzii occurrence in Venezuela. MAP 134. Crocodeilanthe stergiosii occurrence in Venezuela.
NUMBER 100 • 135

Cyrtochilum Kunth

Cyrtochilum Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 280. 1815 [1816]; 

ibid. [qu.] 1: 349. 1815 [1816]. Oncidium sect. Cyrtochilum (Kunth) 

Lindl., Sert. Orchid. t. 48. 1842. Oncidium subgen. Cyrtochilum

(Kunth) Garay, Taxon 19: 444. 1970.

Neodryas Rchb. f., Bot. Zeitung (Berlin) 10: 834. 1852.

Rusbyella Rolfe ex Rusby, Mem. Torrey Bot. Club 6: 122. 1896.

Buesiella C. Schweinf., Bot. Mus. Leafl. 15: 153. 1952.

Dasyglossum Königer & Schildh., Arcula 1: 5. 1994.

Trigonochilum Königer & Schildh., Arcula 1: 13. 1994.

Siederella Szlach. et al., Biodivers. Res. Conservation 1–2: 4. 2006.

Irenea Szlach. et al., Biodivers. Res. Conservation 1–2: 5. 2006.

Epiphytic, lithophytic or terrestrial herbs; caespitose or longcreeping; usually medium to very large in size (compared to other 

Oncidiinae), rarely very small. Roots thick (much thicker than 

those of Oncidium Sw. and Odontoglossum Kunth), long, white. 

Rhizomes thick, conspicuous, usually elongate with pseudobulbs 

well separated from each other, in other cases shorter, loosely covered with scarious, tubular sheaths. Pseudobulbs generally large, 

subhemispherical to ovoid, apically (1-) 2- or 3- (4)-foliate, covered by several distichous sheaths, of which the innermost 2–4 

have foliar blades, surface opaque. Inflorescences 1 or 2, lateral, 

usually arising from sheaths of new buds or from between sheaths 

of mature pseudobulbs, usually paniculate, less commonly racemose, many-flowered, short and ± rigidly erect to very long (to 

several meters long) and creeping, twining or pendulous; peduncles terete (rarely 2- or 3-costate), relatively thick, loosely covered 

with tubular sheaths, sheaths subtending inflorescence branches 

usually conspicuous; floral bracts small or medium-sized, always 

much shorter than the pedicellate ovary, loosely tubular to patent or inflated, ovate, acute. Flowers resupinate, small to very 

large and conspicuous, spreading, lasting several days, variously 

colored, white, yellow to chocolate-brown or purple with lines 

or areas of purple, brown or red, often the petals and sepals different colors, fleshy; perianth segments often abruptly attenuate 

basally and with a conspicuous claw. Sepals free or lateral sepals 

fused below the middle, subequal. Petals free, wider or narrower 

than sepals or differently shaped. Labellum usually narrower 

and fleshier than other perianth segments, simple or somewhat 

3-lobed, usually triangular in general outline or ovate-triangular 

and acute, less commonly pandurate and rounded to truncate apically, the upper 1⁄2 of the labellum often reflexed (in living material); disk with 1 to several calli, these simple or very complex, 

the labellar blade sessile at the base of the column or with a short 

hook; column proportionally long, subterete, making an angle of 

~80°–120° with respect to the labellar blade or less commonly 

the basal part completely parallel to the labellum, straight or sigmoid, often attenuate basally, apically with or without a pair of 

small wings or teeth on each side of the stigma; stigma ventral, 

entire, subquadrate, cordate or transversely elliptic; rostellum 

small, transverse; anther apical, incumbent, operculate; pollinia 

2, ellipsoid or obovoid, waxy, yellow, on a short and wide tegula; 

viscidium small, rounded, ventrally flattened, with relatively conspicuous caudicles. Ovary glabrous. Capsules triangular. [Epidendroideae: Cymbidieae: Oncidiinae.]

A neotropical genus of 115–130 species found in the West 

Indies and South America (Colombia, Venezuela, Ecuador, Peru, 

and Bolivia). Only 1 species appears to be native to the West 

Indies and 1 South American species is adventive in the Greater 

Antilles. The genus is most diverse in the Andes of Colombia and 

Ecuador; ~20 species occur in Venezuela.

References. Chase (1997, 2002); Dalström (2001); 

Garay (1970); Pridgeon et al. (2009); Senghas (1997b); Williams 

et al. (2001).

Species of Cyrtochilum can be found in the shade but are 

more commonly sun loving.

As noted by Chase in Pridgeon et al. (2009: 509), “the taxonomic history of Cyrtochilum is long and troubled.” The species currently assigned to this genus were in the past included in 

several other genera, including Oncidium and Odontoglossum, 

with which they shared evolutionarily convergent floral characters. Cyrtochilum, as conceived here and circumscribed by both 

morphological and molecular characters, is mostly a high Andean group characterized by large, usually long-creeping plants, 

with thick white roots and suborbicular or ovoid pseudobulbs, 

which are not or slightly compressed laterally, and covered with 

various leaf sheaths with foliar blades. The inflorescences are 

usually elongate, often creeping or twining. The flowers are variable in size and include some of the largest in the subtribe and 

in the family (e.g., C. macranthum (Lindl.) Kraenzl.) and usually 

have relatively small and triangular labella. The angle formed by 

the base of the labellum and the column was used in the past to 

determine whether or not a species was assigned to Oncidium

(labellum perpendicular to the column) or Odontoglossum (labellum basally parallel to the column). The “true” members of 

these two genera have, among other differences, strongly laterally compressed pseudobulbs and thin roots.

KEY TO THE SPECIES OF CYRTOCHILUM

1a. Dorsal sepal <1.5 cm long; pedicels at least 2× longer than the sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Rhizomes long-creeping; pseudobulbs separate on the rhizomes; flowers campanulate (in living material), secund; 

pedicels curved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. megalophium

2b. Rhizomes abbreviated or shortly creeping; pseudobulbs aggregated; flowers spreading, not secund; pedicels straight 

(or if curved, then not secund) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Lateral sepals fused in their basal 1⁄2; labellum ± the same width as petals . . . . . . . . . . . . . . . . . . . . . C. distans

 3b. Lateral sepals completely free; labellum conspicuously wider than petals . . . . . . . . . . . . . . . . . C. cimiciferum
136 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

1b. Dorsal sepal ≥2.5 cm; pedicels up to 1.5× longer than the sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4a. Rhizomes short with pseudobulbs aggregated (sometimes only 1 pseudobulb on an herbarium specimen, but often 

>1); inflorescences rigidly erect, emerging from the developing pseudobulb; labellum ± as wide as sepals; callus with 

ridges and linear teeth; column without wings or teeth; flowers white with pink . . . . . . . . . . . . . C. ramosissimum

4b. Rhizomes long-creeping with separate pseudobulbs (almost always only 1 pseudobulb on an herbarium specimen); 

inflorescences creeping, prostrate or twining, several meters long, emerging from mature pseudobulbs (or from 

young ones in C. orgyale); labellum always conspicuously narrower than sepals; callus with ridges and lamellar teeth; 

column with teeth on the sides of the stigma; flowers of various colors but if white, then with areas of dark purple 

or bright red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 5a. Petals triangular, <2× as long as wide; inflorescences emerging from developing pseudobulbs (or from new buds); 

petals pale purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. orgyale

 5b. Petals elliptic or triangular, at least 2.5× as long as wide; inflorescences emerging from mature pseudobulbs; 

petals white with purple areas or brown with yellow areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 6a. Flowers brown or reddish-brown with yellow areas toward the apices of the perianth segments; dorsal sepal 

conspicuously wider than petals that are curved (in living material); labellum panduriform, ± the same width 

from base to apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. falcipetalum

 6b. Flowers white with dark purple or red areas; dorsal sepal the same width to slightly wider than petals; labellum triangular, conspicuously wider toward base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 7a. Sepals ~2 cm long; callus occupying only the central area of the labellar disk . . . . . . . . . C. zebrinum

 7b. Sepals ~3 cm long; callus occupying the entire labellar disk . . . . . . . . . . . . . . . . . . . . . . . . C. detortum

Cyrtochilum cimiciferum (Rchb. f.) Dalström, Lindleyana 16: 

61. 2001; Fernández, Orquídeas Nat. Táchira 50. 2003; 

Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 1: 116–117, fig. 14. 2011. Odontoglossum cimiciferum Rchb. f., Linnaea 22: 849. 1849. Oncidium cimiciferum (Rchb. f.) Beer, Prakt. Stud. Orchid. 274. 1854; 

Dunsterville and Garay, Venez. Orchids Ill. 4: 186–187. 

1966; Foldats, Fl. Venez. 15(5): 300–303, fig. 867, excluding A. 1970; Ortega et al., BioLlania 5: 48. 1987; Romero 

and Carnevali, Orchids Venez., 2nd ed., 2: 676. 2000; Dorr 

et al., Contr. U.S. Natl. Herb. 40: 52. 2000 [2001].

Cyrtochilum flexuosum Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 280. 1815 

[1816]; ibid. [qu.] 1: 350. 1815 [1816].

Terrestrial or epiphytic herbs. Leaf blades 23–50 cm long. 

Flowers 1.5–2 cm in diameter. Perianth pale yellow with opaque, 

reddish-brown spots. Dorsal sepal 7–9 mm long. Callus bright 

yellow.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). In Venezuela, known from the Andes (Mérida, 

Táchira, and Trujillo) and the Sierra de Perijá (Zulia; Map 136). 

In the park, found along the Boconó–Guaramacal road; 2,250 m.

Cyrtochilum detortum (Rchb. f.) Kraenzl., Notizbl. Bot. Gart. 

Berlin-Dahlem 7: 92. 1917; Fernández, Orquídeas Nat. 

Táchira 52. 2003. Oncidium detortum Rchb. f., Gard. 

Chron., ser. 3, 3: 392. 1888; Romero and Carnevali, Orchids Venez., 2nd ed., 2: 680. 2000; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 52. 2000 [2001].

Large terrestrial or rarely epiphytic herbs. Leaf blades 

30–60 cm long. Inflorescences creeping, pendulous or twining, 

3–6 m long. Flowers 4–5 cm in diameter. Perianth white with red 

or reddish-purple spots. Dorsal sepal 2.5–3 cm long.

Found in South America (Venezuela, Ecuador, and Peru). In 

Venezuela, known only from the Andes (Táchira and Trujillo; Map 

137). Near the park boundary in the Laguna de Aguas Negras; 

~1,850 m.

Cyrtochilum distans (Rchb. f.) Kraenzl., Notizbl. Bot. Gart. 

Berlin-Dahlem 7: 99. 1917. Odontoglossum distans Rchb. f., 

Linnaea 22: 848. 1849; Dunsterville and Garay, Venez. Orchids Ill. 4: 170–171. 1966; Foldats, Fl. Venez. 15(5): 210–

MAP 136. Cyrtochilum cimiciferum occurrence in Venezuela.
NUMBER 100 • 137

212, fig. 839. 1970. Symphyglossum distans (Rchb. f.) Garay 

& Dunst., in Dunsterville and Garay, Venez. Orchids Ill. 6: 

40. 1976; Ortega et al., BioLlania 5: 50. 1987; Romero and 

Carnevali, Orchids Venez., 2nd ed., 3: 1065. 2000; Dorr et 

al., Contr. U.S. Natl. Herb. 40: 54. 2000 [2001]; Fernández, 

Orquídeas Nat. Táchira 227. 2003. Trigonochilum distans

(Rchb. f.) Senghas, Orchideen I/C, 33–36: 2211. 1997.

Epiphytic or subterrestrial herbs; pseudobulbs 5–10 cm 

long. Leaf blades to 50 cm long. Inflorescences to 1 m long. 

Flowers 1.5–2 cm in diameter. Perianth pale yellowish-green or 

white with a pink labellum, white basally. Dorsal sepal 9–13 mm 

long. Labellum 7–10 mm long.

Found in South America (Colombia, Venezuela, and Ecuador). In Venezuela, found in the Andes (Mérida, Táchira, and 

Trujillo) and in the Cordillera de la Costa (Aragua and Distrito 

Federal; Map 138). In the park, found in cloud forest along the 

Boconó–Guaramacal road; 2,100–2,450 m.

Cyrtochilum falcipetalum (Lindl.) Kraenzl., Notizbl. Bot. Gart. 

Berlin-Dahlem 7: 92. 1917; Fernández, Orquídeas Nat. 

Táchira 52. 2003; Morillo, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 1: 119–121, fig. 17. 2011. Oncidium falcipetalum Lindl., Orchid. Linden. 14. 1846; Dunsterville and Garay, Venez. Orchids Ill. 1: 266–267. 1959; 

Foldats, Fl. Venez. 15(5): 320–323, fig. 874. 1970; Ortega et 

al., BioLlania 5: 48. 1987; Romero and Carnevali, Orchids 

Venez., 2nd ed., 2: 683. 2000; Dorr et al., Contr. U.S. Natl. 

Herb. 40: 52. 2000 [2001].

Epiphytic or terrestrial herbs; pseudobulbs 4–15 cm long. 

Leaf blades 10–60 cm long. Inflorescences 2–4 m long. Flowers 

brown or reddish-brown. Sepals 3.5–4.5 cm long. Petals and labellum with yellow spots.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). In Venezuela, known from the Andes (Lara, 

Mérida, Táchira, and Trujillo) and the Cordillera de la Costa 

(Aragua and Distrito Federal; Map 139). Park records are from 

both slopes of Guaramacal; 1,950–2,350(–3,000) m.

MAP 138. Cyrtochilum distans occurrence in Venezuela. MAP 139. Cyrtochilum falcipetalum occurrence in Venezuela.

MAP 137. Cyrtochilum detortum occurrence in Venezuela.
138 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Cyrtochilum megalophium (Lindl.) Kraenzl., Notizbl. Bot. 

Gart. Berlin-Dahlem 7: 98. 1917; Fernández, Orquídeas 

Nat. Táchira 54. 2003; Morillo, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 1: 122, fig. 182. 2011. Odontoglossum megalophium Lindl., Orchid. Linden. 15. 1846; 

Dunsterville and Garay, Venez. Orchids Ill. 3: 210–211. 

1965; Foldats, Fl. Venez. 15(5): 221–223, fig. 844. 1970; 

Ortega et al., BioLlania 5: 48. 1987; Romero and Carnevali, 

Orchids Venez., 2nd ed., 2: 661. 2000; Dorr et al., Contr. 

U.S. Natl. Herb. 40: 52. 2000 [2001].

Epiphytic or terrestrial herbs; pseudobulbs 4–10 × 1.5–4 

cm. Leaf blades 18–55 cm long. Inflorescences broadly paniculate, suberect to twining, to 2 m long. Flowers yellow. Dorsal 

sepal 4–6 mm long; lateral sepals 7–8 mm long.

Found in South America (Colombia, Venezuela, and Peru). 

In Venezuela, restricted to the Andes (Mérida, Táchira, and Trujillo; Map 140). In the park, found in páramo on the Fila de Los 

Recostaderos and in cloud forest on the north slope of Guaramacal; 2,700–3,100 m.

Cyrtochilum orgyale (Rchb. f. & Warsz.) Kraenzl., Notizbl. Bot. 

Gart. Berlin-Dahlem 7: 92. 1917; Fernández, Orquídeas 

Nat. Táchira 55. 2003. Oncidium orgyale Rchb. f. & 

Warsz., Bonplandia (Hannover) 2: 102. 1854; Foldats, Fl. 

Venez. 15(5): 370–373, fig. 891. 1970; Ortega et al., BioLlania 5: 48. 1987; Romero and Carnevali, Orchids Venez., 

2nd ed., 2: 694. 2000; Dorr et al., Contr. U.S. Natl. Herb. 

40: 52. 2000 [2001].

Cyrtochilum undulatum auct., non Kunth; Morillo, in Morillo et al., Bot. 

Ecol. Monocot. Páramos Venez. 1: 126–127, fig. 21, foto 11. 2011.

Large, epiphytic herbs; pseudobulbs to 15 cm long. Leaf 

blades 15–50 cm long. Inflorescences 3–4 m long. Sepals 

brownish-purple; dorsal sepal 2.4–2.7 cm long. Petals purple.

Found in South America (Colombia and Venezuela). In 

Venezuela, restricted to the Andes (Mérida, Táchira, and Trujillo; Map 141). In the park, found on the north slope and in the 

Páramo de Guaramacal; 2,250–3,000 m.

Dalström (2001: 71) speculated that Cyrtochilum orgyale

might be synonymous with C. undulatum Kunth. If this proves 

to be true, then the latter name has priority.

Cyrtochilum ramosissimum (Lindl.) Dalström, Lindleyana 16: 

72. 2001; Morillo, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 1: 123–124, fig. 19, foto 10. 2011. Odontoglossum ramosissimum Lindl., Fol. Orchid. 1: Odontoglossum 16. 1852; Dunsterville and Garay, Venez. Orchids Ill. 

3: 212–213. 1965; Foldats, Fl. Venez. 15(5): 229–231, fig. 

847. 1970; Ortega et al., BioLlania 5: 48. 1987; Romero 

and Carnevali, Orchids Venez., 2nd ed., 2: 664. 2000; Dorr 

et al., Contr. U.S. Natl. Herb. 40: 52. 2000 [2001].

Epiphytic herbs; pseudobulbs 6.5–12 × 3–5 cm. Leaf blades 

30–60 cm long. Inflorescences 60–120 cm tall, erect. Flowers 

white, bases of perianth segments purple. Sepals 2.5–3 cm long.

Found in South America (Colombia, Venezuela, Ecuador, and Peru). In Venezuela, restricted to the Andes (Mérida, 

Táchira, and Trujillo; Map 142). In the park, found in cloud forest along the Boconó–Guaramacal road; 1,850–2,150 m.

Cyrtochilum zebrinum (Rchb. f.) Kraenzl., Notizbl. Bot. Gart. 

Berlin-Dahlem 7: 94. 1917; Fernández, Orquídeas Nat. 

MAP 140. Cyrtochilum megalophium occurrence in Venezuela. MAP 141. Cyrtochilum orgyale occurrence in Venezuela.
NUMBER 100 • 139

Táchira 57. 2003; Morillo, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 1: 127, fig. 22. 2011. Odontoglossum zebrinum Rchb. f., Linnaea 22: 849. 1849; Dunsterville 

and Garay, Venez. Orchids Ill. 2: 260–261. 1961. Oncidium 

zebrinum (Rchb. f.) Rchb. f., Bonplandia (Hannover) 2: 12. 

1854; Foldats, Fl. Venez. 15(5): 404–406, fig. 901. 1970; 

Dorr et al., Contr. U.S. Natl. Herb. 40: 52. 2000 [2001].

Oncidium engelii Rchb. f., Linnaea 41: 22. 1877 [1876]; Dunsterville and 

Garay, Venez. Orchids Ill. 4: 189–190. 1966; Foldats, Fl. Venez. 15(5): 

316–317. 1970; Romero and Carnevali, Orchids Venez., 2nd ed., 2: 

681. 2000.

Oncidium detortum auct., non Rchb. f.; Dunsterville and Garay, Venez. Orchids Ill. 3: 222–223. 1965.

Epiphytic or more frequently terrestrial herbs; pseudobulbs 

large, to 10 cm long. Leaf blades 15–60 cm long. Inflorescences 

3–4 m long, creeping or twining. Perianth white with brown or 

red bands. Dorsal sepal ~2 cm long.

Found in South America (Venezuela, Colombia, Ecuador, 

and Peru). In Venezuela, found in the Andes (Lara, Mérida, Portuguesa, Táchira, and Trujillo) and in the Cordillera de la Costa 

(Distrito Federal and Miranda; Map 143). In the park, found on 

the Fila de Agua Fría and both slopes of Guaramacal; 1,800–

2,800(–3,000) m.

Dichaea Lindl.

Dichaea Lindl., Gen. Sp. Orchid. Pl. 208. 1833.

Epiphytic, rarely lithophytic or subterrestrial herbs; pendent, 

arched, erect or prostrate; caespitose; pseudobulbs absent. Roots 

usually thin, sometimes very thick and fleshy, arising from the 

bases of stems or in prostrate plants internodes in contact with 

the substrate. Stems monopodial or pseudomonopodial with 

indeterminate growth, branching from the base (and creating 

caespitose plants) or less commonly from more distal internodes, 

completely obscured by leaf sheaths that are persistent and only 

disintegrate to reveal terete stems on very old plants. Leaves distichous, articulate or marcescent; sheaths imbricate; leaf blades 

in the same plane as sheaths or perpendicular to them, usually 

oblong or linear-oblong, sometimes elliptic or narrowly ovateelliptic, rounded to acute or acuminate, often apiculate or with 

a subapical mucro, margins entire, ciliate to finely toothed, cilia 

or teeth often restricted to the upper part of the blade. Inflorescences 1-flowered, solitary, opposite the leaves; peduncles usually filiform, usually shorter than the leaves; floral bracts usually 

clasping more than 1⁄2 the length of the ovary, tubular, apiculate 

or mucronate. Flowers resupinate, campanulate or the segments ± spreading, fugacious; perianth segments oblong-elliptic 

to ovate (rarely obovate), rounded to acuminate, outer surface 

smooth to verruculose, usually white or green, rarely yellow or 

pink, often spotted or tinged with purple or pink, 1–5-nerved, 

membranous to subfleshy. Lateral sepals somewhat oblique. Petals similar to sepals or narrower. Labellum rigidly attached to 

the column base, usually attenuate basally into a conspicuous 

claw, rarely subsessile, the claws ecallose or rarely with 1 or 2 

calli; labellar blade usually anchor-shaped, less commonly pandurate, margins entire or finely ciliolate, the same color as other 

members of the perianth but often with larger areas of color (as 

described above), usually ecallose; column short to somewhat 

elongate, subcylindric, often forming an angle with the labellum; 

anther terminal, operculate, incumbent, clinandrium shallow; 

MAP 143. Cyrtochilum zebrinum occurrence in Venezuela.

MAP 142. Cyrtochilum ramosissimum occurrence in Venezuela.
140 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

pollinia 4, ovoid, waxy, tegula well developed, short to elongate; 

viscidium small; stigma relatively large, ventral but occasionally 

appearing frontal because of the angle of the column, often with 

an imperceptibly pubescent ligule. Ovary obconical to fusiform, 

smooth to densely muricate, usually with a linear bracteole at 

the base; bracteole longer to shorter than the ovary. Capsules 

ellipsoid, smooth or muricate. [Epidendroideae: Cymbidieae: 

Zygopetalinae.]

A neotropical genus of ~100 species found in Mexico, Central America, the West Indies, and South America (Colombia, 

Venezuela, the Guianas, Ecuador, Peru, Bolivia, and Brazil); ~16 

species occur in Venezuela.

References. Folsom (1996); Neubig et al. (2009); 

Pridgeon et al. (2009); Senghas (1994a).

Molecular and morphological data support the monophyly 

of Dichaea. Molecular data (Neubig et al., 2009), however, call 

into question the infrageneric classification that was constructed 

on morphology alone. In particular, Neubig et al. (2009) suggest 

that D. sect. Dichaea should be expanded to include D. sect. 

Dichaeastrum.

Dichaea has been difficult to study because pressed flowers 

do not rehydrate well, and as a consequence, herbarium material 

alone generally is inadequate for interpreting floral morphology. 

To make the situation even more complicated, plants usually 

produce flowers in succession and have only 1(2) flower open 

per stem. These flowers often remain attached to the paper used 

to press them; as a consequence most herbaria are filled with 

sterile specimens.

KEY TO THE SPECIES OF DICHAEA

1a. Leaves articulate with their sheaths, eventually deciduous and leaving a scar at the apex of the sheath . . . . . . . . . . . . 2

2a. Mature leaf blades oblong to elliptic-oblong or elliptic, 38–63(–75) mm long, apices rounded or subacute and 

mucronate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. robusta

2b. Mature leaf blades oblong-lanceolate, 22–40 mm long, apices acuminate to narrowly acute . . . . D. camaridioides

1b. Leaves not articulate with their sheaths, marcescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Plants small, usually <10 cm long; leaf blades 2.5–3 mm wide, margins conspicuously ciliate-denticulate . . . . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. hystricina

3b. Plants larger, usually >15 cm long; leaf blades 3–7.5 mm wide, margins smooth or minutely denticulate only in the 

apical half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. latifolia

Dichaea camaridioides Schltr., Repert. Spec. Nov. Regni Veg. 

Beih. 7: 201. 1920; Romero and Carnevali, Orchids Venez., 

2nd ed., 1: 152. 2000; Carnevali and Ramírez-Morillo, in 

Berry et al., Fl. Venez. Guayana 7: 301. 2003; Fernández, 

Orquídeas Nat. Táchira 58. 2003.

Dichaea brachypoda auct., non Rchb. f.; Foldats, Fl. Venez. 15(5): 447–449, 

fig. 916. 1970; Dunsterville and Garay, Venez. Orchids Ill. 3: 72–73. 

1965.

Epiphytic herbs; erect or arching to pendulous; stems to 30 

cm long. Leaves articulate with their sheaths; leaf blades oblonglanceolate, 22–40 × 5–10 mm, margins smooth, microscopically 

pilose toward apex, apices acute to acuminate. Flowers creamcolored, purple-speckled, margins green. Sepals 7–13 mm long.

Found in South America (Colombia, Venezuela, and possibly Guyana). In Venezuela, known from the Andes (Lara, Portuguesa, and Táchira), the Cordillera de la Costa (Yaracuy), and 

the Venezuelan Guayana (Amazonas; Map 144). In the park, collected on the south slope of Guaramacal near El Campamento 

of El Santuario and on the Camino Real Paramito–Batatal; 

(1,550−)1,800−2,000 m.

Dichaea hystricina Rchb. f., Flora 48: 279. 1865; Dunsterville 

and Garay, Venez. Orchids Ill. 2: 86–87. 1961; Foldats, Fl. 

Venez. 15(5): 452–454, fig. 918. 1970; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 155. 2000; Carnevali and 

Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 7: 302, 

fig. 290. 2003; Fernández, Orquídeas Nat. Táchira 59. 2003.

Epiphytic herbs; suberect, spreading or pendulous; stems 

branched or not, 3–8(–15) cm long. Leaves not articulate with 

their sheaths; leaf blades elliptic to elliptic-lanceolate, 4–10 

× 1.5–3 mm, margins conspicuously denticulate-ciliate, apices acute to acuminate. Flowers white or greenish-white with 

MAP 144. Dichaea camaridioides occurrence in Venezuela.
NUMBER 100 • 141

purple-pink spots. Sepals 4−6.5 mm long. Labellum occasionally 

completely violet.

Found in Central America, the West Indies, and South America 

(Colombia, Venezuela, Trinidad and Tobago, Ecuador, and Brazil). 

In Venezuela, found in the Andes (Portuguesa and Táchira), the 

Cordillera de la Costa (Anzoátegui, Aragua, Carabobo, Sucre, and 

Yaracuy), and the Venezuelan Guayana (Amazonas and Bolívar; 

Map 145). La Divisoria de la Concepción; 1,700 m.

The diminutive Dichaea hystricina is usually collected in 

shade growing among bryophytes. It is the only Dichaea in our 

flora with conspicuously denticulate-ciliate leaf margins.

Dichaea latifolia Lindl., Gen. Sp. Orchid. Pl. 208. 1833.

Dichaea longa Schltr., Repert. Spec. Nov. Regni Veg. Beih. 10: 54. 1922. 

Dichaea latifolia var. longa (Schltr.) Folsom, Orchid Digest 60: 154. 

1996; Romero and Carnevali, Orchid Venez., 2nd ed., 1: 156. 2000; 

Carnevali and Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 7: 

302. 2003.

Dichaea muricata auct., non (Sw.) Lindl.; Dunsterville and Garay, Venez. 

Orchids Ill. 2: 89. 1961; Foldats, Fl. Venez. 15(5): 460–463, fig. 921. 

1970.

Epiphytic herbs; pendulous; stems unbranched or branched 

only in the lower 1⁄2, to 1 m long. Leaves articulate with their 

sheaths; leaf blades elliptic or rarely elliptic-lanceolate, 8–20 × 

3–7.5 mm, margins smooth or minutely denticulate above, apices obtuse or rounded. Flowers pale greenish-brown, sometimes 

purple-spotted. Sepals 7–14 mm long. Labellum white with violet zones.

Found in the West Indies and South America (Colombia, 

Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, found in 

the Andes (Portuguesa and Táchira), the Cordillera de la Costa 

(Anzoátegui, Aragua, Miranda, Monagas, and Sucre), the Venezuelan Guayana (Bolívar), and Nueva Esparta (Map 146). 

Cloud forest near the lower boundaries of both slopes of the 

park, including the Laguna de Aguas Negras, El Campamento 

below Cerro El Diablo, and La Divisoria de la Concepción; 

(1,300−)1,800−2,000 m.

According to Folsom (1996: 152), Dichaea latifolia var. 

longa is distinguished from the nominate variety by having “leaf 

blades spreading, nearly touching” versus “leaf blades usually 

retrorse, spaced.” We fail to see this distinction in the Venezuelan 

material available to us and have adopted a broader concept of 

D. latifolia.

Dichaea robusta Schltr., Repert. Spec. Nov. Regni Veg. 27: 83. 

1929; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 

160. 2000; Carnevali and Ramírez-Morillo, in Berry et al., 

Fl. Venez. Guayana 7: 302. 2003.

Dichaea morrisii auct., non Fawc. & Rendle; Dunsterville and Garay, Venez. 

Orchids Ill. 1: 94–95. 1959, pro parte; Foldats, Fl. Venez. 15(5): 457–

459, fig. 920. 1970; Dorr et al., Contr. U.S. Natl. Herb. 40: 49. 2000 

[2001].

Epiphytic herbs; erect or pendulous; stems 10–55 cm long. 

Leaves articulate with their sheaths; leaf blades oblong to ellipticoblong or elliptic, 38–63(–75) × 10−16 mm, margins smooth, 

apices rounded or subacute and mucronate. Flowers green or 

yellowish-green, sometimes streaked with brown or purple. Sepals 7−18 mm long. Labellum purple or (less commonly) whitish 

with purple or brown blotches.

Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, Ecuador, Peru, and Bolivia). Venezuelan collections are from the Andes (Mérida, 

MAP 145. Dichaea hystricina occurrence in Venezuela. MAP 146. Dichaea latifolia occurrence in Venezuela.
142 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Portuguesa, and Táchira), the Sierra de Perijá (Zulia), the Cordillera de la Costa (Miranda and Yaracuy), and the Venezuelan 

Guayana (Amazonas, Bolívar, and Delta Amacuro; Map 147). 

Near La Divisoria de la Concepción; 1,700 m.

Dichaea robusta Schltr. is part of a complex of at least two 

species (R. Dressler, pers. comm.), both of which have been confused under one name. One of them, to which the name D. robusta applies, occurs from Costa Rica southward to at least 

Peru. The second entity, D. morissii Fawcett & Rendle, occurs 

in the West Indies (type locality) and Central America. They are 

sympatric in Costa Rica. Dichaea robusta features broader sepals and petals that are acute to obtuse at the apex, the claw 

of the labellum is thick, the lateral lobes of the labellum are 

long and reflexed, and the labellum apex is more or less truncate to rounded. Dichaea morrisii, on the other hand, displays 

narrower, narrowly acute to acuminate sepals and petals, short, 

straight lateral lobes to the labellum, and the labellum apex is 

acute. Flower color is also different. Dichaea robusta usually has 

greenish or green-white flowers with a few red or purple spots, 

whereas D. morrisii is characterized by white flowers with pinkpurple stripes. Both species have dark purple-red labella, these 

somewhat darker in D. robusta.

Elleanthus C. Presl

Elleanthus C. Presl, Reliq. Haenk. 1: 97. 1830 [1827].

Terrestrial, epiphytic or subterrestrial herbs; erect to pendulous; caespitose, often forming dense clumps. Rhizomes short. 

Roots relatively thick, glabrous. Stems thin to relatively thick, 

± woody, elongate, simple or branched, to 2(–3.5) m tall, often 

articulate, terete or ± laterally compressed, covered by sheaths, 

usually leafy in the apical 3⁄4. Leaves convolute, plicate or rarely 

subconduplicate, articulate, distichous or rarely subdistichous, 

soft or rigid, sessile; leaf sheaths amplexicaul to funnel-shaped, 

often pubescent or verruculose; leaf blades linear to lanceolate, nerves prominent, especially in the lower surface, usually 

glabrous. Inflorescences terminal or (in some Andean species) 

originating directly from rhizomes, erect, racemose, usually with 

many spiraled or few distichous flowers, or capitate, always 

shorter than the stem, usually subtended by several foliaceous 

or scarious bracts; floral bracts usually conspicuous and brightly 

colored, rarely inconspicuous or scarious. Flowers resupinate or 

not, usually campanulate, mostly brightly colored, sometimes 

white; perianth segments membranous. Sepals free, subequal, 

mostly elliptic, lanceolate or narrowly ovate. Petals similar to 

sepals but usually narrower and apically more obtuse. Labellum free from column to base, erect, subequal or longer than the 

sepals, blade concave in the basal 1⁄2, funnelform and clasping 

the column, apically simple or inconspicuously 3-lobed, margins 

entire, denticulate or fimbriate; disk basally with 1 or 2 prominent calli; column erect, semiterete, sometimes winged, footless; 

anther terminal, operculate, ± incumbent or erect, 2-locular 

or imperfectly 4-locular; clinandrium small; pollinia 8, waxy 

or soft, ovoid, without viscidium; rostellum transverse, stigma 

ventral, emergent. Ovary relatively thick and short, smooth or 

verruculose, pedicellate. Capsules cylindrical. [Epidendroideae: 

Sobralieae.]

A neotropical genus of ~100 species found in Mexico, Central America, the West Indies, and South America (Colombia, 

Venezuela, Guyana, Suriname, Ecuador, Peru, Bolivia, and Brazil); most diverse in the Andes. Approximately 20 species occur 

in Venezuela.

References. Løjtnant (1977); Neubig et al. (2011); 

Pridgeon et al. (2005).

Molecular data support the monophyly of Elleanthus (Neubig et al., 2011). Most members of the genus have brightly colored bracts and flowers, and most species are assumed to be 

pollinated by hummingbirds.

KEY TO THE SPECIES OF ELLEANTHUS

1a. Plants always epiphytic; leaves linear-oblong, <7 mm wide; inflorescences laterally compressed with distichous, imbricate 

floral bracts; flowers white, barely emerging from the condensed inflorescence, inconspicuous; sepals 4–6 mm long . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. graminifolius

1b. Plants epiphytic or terrestrial; leaves elliptic to lanceolate, >10 mm wide; inflorescences usually racemose or conical, not 

laterally compressed; flowers variously colored but usually not white, conspicuous; sepals >9 mm long . . . . . . . . . . . 2

2a. Plants epiphytic; inflorescences 2–4-flowered, very lax, distichous; flowers arranged in a zigzag pattern . . . . E. confusus

MAP 147. Dichaea robusta occurrence in Venezuela.
NUMBER 100 • 143

2b. Plants terrestrial or epiphytic; inflorescences usually multiflowered, dense or sublax; flowers usually spirally arranged, 

never in a zigzag pattern . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Plants terrestrial, composed of elongated stems that freely produce lateral branches, often fasciculate, 5–10 cm 

long each, at several distances from the base of the main stem; flowers bright orange and yellow or rarely entirely 

orange or yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Flowers bright orange with yellow or entirely orange; lateral sepals lanceolate, acute . . . . . E. aurantiacus

 4b. Flowers pale yellow; lateral sepals triangular-elliptic, acuminate . . . . . . . . . . . . . . . . . . . . . . E. flavescens

 3b. Plants terrestrial or epiphytic, composed of simple, unbranched (rarely branched in E. furfuraceus) stems; flowers 

pinkish or purplish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 5a. Floral bracts at base of inflorescences narrowly elliptic, long-acuminate or setose-acuminate, 2.5–6.5 cm 

long, 2 or 3× longer than the flowers they subtend; dorsal sepal 15–18 mm long . . . . . . . . . . . E. lupulinus

 5b. Floral bracts at base of inflorescences elliptic or ovate, obtuse, acute or short-acuminate, <2.5 cm long, as 

long as or slightly longer than the flowers they subtend; dorsal sepal 5–13 mm long . . . . . . . . . . . . . . . . 6

 6a. Inflorescences with 10 or fewer flowers; stems 1.5–4 mm in diameter, usually <50 cm long (when fertile); 

leaves narrowly lanceolate, 1–2.5 cm wide, with 2 or 3 nerves each side of the midnerve; floral bracts 

4–12 mm wide; dorsal sepal 5–8 mm long; lateral sepals strongly keeled dorsally; basal sacciform portion of labellum accounting for about 1⁄2 of the total labellar length; ovary furfuraceous . . . . . . . . . . 7

 7a. Petals elliptic; labellum deeply sacciform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. furfuraceus

 7b. Petals oblong or oblong-spathulate; labellum shallowly sacciform . . . . . . . . . . . . . . . . E. gracilis

 6b. Inflorescences with 20 or more flowers; stems 5–10 mm in diameter, usually >80 cm long (when fertile); 

leaves lanceolate or oblong-lanceolate, 2.5–7 cm wide, with 4–6 nerves on each side of the midnerve; 

floral bracts 2–6 mm wide; dorsal sepal 9.5–13 mm long; lateral sepals not keeled dorsally; basal sacciform portion of labellum accounting for 1⁄3 or less of total labellar length; ovary glabrous . . . . . . . 8

 8a. Calli at base of labellum discrete, free from each other; unconstricted portion of labellum about as 

long as wide or wider than long; floral bracts 3–6 mm wide . . . . . . . . . . . . . . . . . . E. columnaris

 8b. Calli at base of labellum fused into a ± 2-lobed single structure; unconstricted portion of labellum 

longer than wide; floral bracts to 2 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. wageneri

Elleanthus aurantiacus (Lindl.) Rchb. f., Ann. Bot. Syst. 6: 482. 

1861 [1863]; Dunsterville and Garay, Venez. Orchids Ill. 4: 

68–69. 1966; Foldats, Fl. Venez. 15(1): 211–213, fig. 77. 

1969, pro parte; Romero and Carnevali, Orchids Venez., 

2nd ed., 1: 175. 2000; Morillo, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 1: 129, fig. 23, foto 10. 2011. 

Evelyna aurantiaca Lindl., in Bentham, Pl. Hartw. 149. 

1844 [1845].

Elleanthus gracilis auct., non (Rchb. f.) Rchb. f.; Ortega et al., BioLlania 5: 

47. 1987.

Terrestrial or lithophytic herbs, to 1.5 m tall. Leaf blades 

2–12 × 0.5–2.4 cm. Flowers usually orange and yellow. Dorsal 

sepal 4.5–8.2 mm long.

Found in Central America and South America (Colombia, 

Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, known 

from the Andes (Mérida, Táchira, and Trujillo) and the Cordillera de la Costa (Aragua, Distrito Federal, Miranda, and Yaracuy; Map 148). Collected in cloud forest in the park without 

precise locality; 2,000–2,800 m.

Elleanthus aurantiacus is a species of wide distribution that 

frequently forms large colonies in forest clearings or on road cuts. 

The label on one collection from the park (Stergios et al. 6536) 

describes the flowers as “azul-moradas,” which probably can 

be attributed to the specimen being collected postanthesis since 

floral tissues turn black with age. We treat a yellow-flowered 

variant in the E. aurantiacus complex as a distinct species, E. flavescens (Lindl.) Rchb. f.

Elleanthus columnaris (Lindl.) Rchb. f., Ann. Bot. Syst. 6: 483. 

1861 [1863]; Dunsterville and Garay, Venez. Orchids Ill. 2: 

MAP 148. Elleanthus aurantiacus occurrence in Venezuela.
144 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

94–95. 1961; Foldats, Fl. Venez. 15(1): 217–219, fig. 80. 

1969; Romero and Carnevali, Orchids Venez., 2nd ed., 

1: 177. 2000; Carnevali and Ramírez-Morillo, in Berry et 

al., Fl. Venez. Guayana 7: 313, fig. 298. 2003; Fernández, 

Orquídeas Nat. Táchira 62. 2003. Evelyna columnaris

Lindl., Orchid. Linden. 11. 1846.

Terrestrial, lithophytic or epiphytic herbs, 0.6–1.5 m tall. 

Leaf blades 1.3–3 × 3–6.5 cm. Bracts and flowers bright purple 

or magenta at anthesis. Dorsal sepal 9–13 mm long. Petals white 

with pink.

Found in South America (Colombia, Venezuela, Ecuador, Bolivia, and Brazil). In Venezuela, found in Amazonas, Bolívar, Falcón, 

Lara, Mérida, Táchira, and Trujillo (Map 149). In the park, found 

in cloud forest on both slopes of Guaramacal; 1,700–2,400 m.

Elleanthus confusus Garay, Bot. Mus. Leafl. 26: 13. 1978; Carnevali and Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 7: 313, fig. 298. 2003.

Elleanthus kermesinus auct., non (Lindl.) Rchb. f.; Foldats, Fl. Venez. 15(1): 

226, fig. 83. 1970.

Elleanthus virgatus auct., non (Rchb. f.) C. Schweinf.; Dunsterville and Garay, Venez. Orchids Ill. 2: 100–101. 1961.

Epiphytic herbs, erect to pendulous, straggling. Leaf blades 

3–9 × 0.5–1.5 cm. Bracts and flowers pink-lavender to deep purple. Dorsal sepal 8–10 mm long.

Found in South America (Venezuela and possibly adjacent 

Brazil). In Venezuela, restricted to high elevations in the Andes 

(Táchira and Trujillo), the Sierra de Perijá (Zulia), the Cordillera 

de la Costa (Aragua), and the Venezuelan Guayana (Amazonas 

and Bolívar; Map 150). In the park, collected once in forest on 

the south slope; 1,850 m.

Elleanthus flavescens (Lindl.) Rchb. f., Ann. Bot. Syst. 6: 479. 

1861 [1862]; Romero and Carnevali, Orchids Venez., 2nd 

ed., 1: 179. 2000. Evelyna flavescens Lindl., Orchid. Linden. 11. 1846.

Terrestrial herbs, erect, 50–80 cm tall. Flowers yellow or pale 

yellow. Leaf blades 5–10 × 0.3–2 cm. Dorsal sepal 5–6 mm long.

Found in South America (Venezuela, Ecuador, and Peru). 

In Venezuela, restricted to the Andes (Mérida, Táchira, and 

MAP 151. Elleanthus flavescens occurrence in Venezuela.

MAP 149. Elleanthus columnaris occurrence in Venezuela. MAP 150. Elleanthus confusus occurrence in Venezuela.
NUMBER 100 • 145

Trujillo) and the Sierra de Perijá (Zulia; Map 151). Collected in 

cloud forest on both slopes of Guaramacal and in the Páramo de 

Guaramacal; (2,000–)2,700–3,100 m.

This species is vegetatively indistinguishable from the much 

more common Elleanthus aurantiacus, and the two are sometimes considered to be conspecific. We choose to recognize them 

as distinct species because the color differences are consistent 

and E. flavescens grows in pure populations.

Elleanthus furfuraceus (Lindl.) Rchb. f., Ann. Bot. Syst. 6: 480. 

1861 [1862]; Dunsterville and Garay, Venez. Orchids Ill. 1: 

104–105. 1959; Foldats, Fl. Venez. 15(1): 221–224, fig. 81. 

1969; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 

180. 2000; Morillo, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 1: 130–131, fig. 24. 2011. Evelyna furfuracea Lindl., Orchid. Linden. 12. 1846.

Epiphytic or more rarely terrestrial herbs, 30–110 cm tall. 

Leaf blades 6–14 × 1–2.5 cm. Flowers bright pink, purple-pink 

or reddish. Dorsal sepal 5–8 mm long.

Found in South America (Colombia, Venezuela, Guyana, 

Ecuador, Peru, and Brazil). Widespread in Venezuela (Amazonas, Aragua, Distrito Federal, Lara, Mérida, Miranda, Monagas, 

Sucre, Táchira, and Trujillo; Map 152). Collected in cloud forest 

on the north slope of Guaramacal; 1,800–2,350(–3,000) m.

Elleanthus gracilis (Rchb. f.) Rchb. f., Ann. Bot. Syst. 6: 481. 

1861 [1863]; Foldats, Fl. Venez. 15(1): 225–226, fig. 82. 

1969; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 

181. 2000. Evelyna gracilis Rchb. f., Linnaea 22: 843. 1849.

Elleanthus kermesinus auct., non (Lindl.) Rchb. f.; Dunsterville and Garay, 

Venez. Orchids Ill. 3: 78–79. 1965.

Epiphytic herbs, erect to pendulous, 20–50 cm tall. Leaf 

blades 7–13 × 0.8–1.3 cm. Bracts and flowers pink, rose-pink or 

pale purple. Dorsal sepal 6.2–7 mm long.

Restricted to South America (Colombia, Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, found in the Cordillera de 

Mérida (Mérida, Portuguesa, and Trujillo) and in the Venezuelan 

Guayana (Amazonas and Bolívar; Map 153). Our collections are 

from fire-induced páramo on the north slope and cloud forest on 

the south slope of Guaramacal; 1,950–2,600 m.

This species is very similar to Elleanthus furfuraceus but is 

distinguished by its narrower floral bracts (1.5–3.5 vs. 4–12 mm 

wide) and petals (1.1–2 vs. 1.7–3.5 mm wide).

Elleanthus graminifolius (Barb. Rodr.) Løjtnant, Bot. Not. 

129: 447. 1977; Romero and Carnevali, Orchid Venez., 

2nd ed., 1: 182. 2000; Carnevali and Ramírez-Morillo, in 

Berry et al., Fl. Venez. Guayana 7: 313–314. 2003; Fernández, Orquídeas Nat. Táchira 63. 2003. Adeneleuterophora 

graminifolia Barb. Rodr., Gen. Spec. Orchid. 2: 171. 1881.

Elleanthus pusillus Schltr., Notizbl. Bot. Gart. Berlin-Dahlem 8: 117. 1922.

Elleanthus linifolius auct., non C. Presl; Dunsterville and Garay, Venez. Orchids Ill. 2: 98–99. 1961; Foldats, Fl. Venez. 15(1): 228–231, fig. 84. 

1969.

Epiphytic herbs, usually erect, sometimes ascending, 10–50 

cm tall. Leaf blades 3–15 × 0.1–0.4 cm. Flowers cream-colored 

or white. Dorsal sepal 3–3.8 mm long.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, 

and Brazil). Widespread in Venezuela (Amazonas, Anzoátegui, 

Aragua, Barinas, Bolívar, Carabobo, Miranda, Portuguesa, and 

Táchira; Map 154). La Divisoria de la Concepción; ~1,500 m.

MAP 153. Elleanthus gracilis occurrence in Venezuela.

MAP 152. Elleanthus furfuraceus occurrence in Venezuela.
146 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Mexican and Central American populations referred to Elleanthus graminifolius may represent a distinct species.

Elleanthus lupulinus (Lindl.) Rchb. f., Ann. Bot. Syst. 6: 483. 

1861 [1863]; Foldats, Fl. Venez. 15(1): 231–233, fig. 85. 

1969; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 

183. 2000; Fernández, Orquídeas Nat. Táchira 63. 2003; 

Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 1: 132, fig. 25, foto 13. 2011. Evelyna lupulina

Lindl., Orchid. Linden. 11. 1846.

Robust, terrestrial herbs, to 1.5 m tall. Leaf blades 15–19 × 

1.7–5.3 cm. Floral bracts magenta. Flowers pinkish-purple. Dorsal sepal 15–18 mm long.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, recorded from the Andes (Mérida, 

Táchira, and Trujillo), the Cordillera de la Costa (Distrito Federal), and the Venezuelan Guayana (Bolívar; Map 155). Cloud 

forest and páramo near the television antennas in the Páramo de 

Guaramacal; 2,900–3,100 m.

This is a robust, showy species that is easy to recognize by its 

long, acuminate floral bracts. Elleanthus lupulinus is sometimes 

considered to be a synonym of E. maculatus (Lindl.) Rchb. f., 

but we follow Garay (1978) in recognizing two distinct species.

Material from Peru identified as Elleanthus lupulinus by 

Bennett and Christenson (1995: t. 233) is less robust with much 

smaller flowers displaying a differently shaped column, and it 

may represent a distinct species.

Elleanthus wageneri (Rchb. f.) Rchb. f., Ann. Bot. Syst. 6: 474. 

1861 [1862]; Dunsterville and Garay, Venez. Orchids Ill. 3: 

80–81. 1965; Foldats, Fl. Venez. 15(1): 234–236, fig. 87. 

1969; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 

189. 2000; Carnevali and Ramírez-Morillo, in Berry et al., Fl. 

Venez. Guayana 7: 315, fig. 299. 2003; Fernández, Orquídeas 

Nat. Táchira 64. 2003; Morillo, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 1: 133, fig. 26, foto 14. 2011. Evelyna wageneri Rchb. f., Bonplandia (Hannover) 2: 21. 1854.

FIGURE 25F

Elleanthus columnaris auct., non (Lindl.) Rchb. f.; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 49. 2000 [2001].

MAP 155. Elleanthus lupulinus occurrence in Venezuela. MAP 156. Elleanthus wageneri occurrence in Venezuela.

MAP 154. Elleanthus graminifolius occurrence in Venezuela.
NUMBER 100 • 147

Terrestrial or epiphytic herbs, 0.5–1.8 m tall. Leaf blades 

12–25 × 2.5–7 cm. Floral bracts bright magenta. Flowers 

maroonish-brown, magenta at base. Dorsal sepal 9.5–11 mm 

long. Petals white flushed with magenta; labellum white with 

brown marks inside.

Known only from Venezuela, where it is found in the Andes 

(Mérida, Táchira, and Trujillo), the Cordillera de la Costa (Distrito Federal), and the Venezuelan Guayana (Bolívar) (Map 156). 

Park records are from cloud forest in Qda. Segovia and the area 

between El Campamento and Cerro El Diablo; 1,950–2,350 m.

This is a showy species superficially similar to Elleanthus 

columnaris but readily distinguished by its narrower floral bracts 

and fused labellar calli. Elleanthus wageneri also can be distinguished by its size as it has the largest leaves of all the Elleanthus

species found in Guaramacal.

Epidendrum L.

Epidendrum L., Sp. Pl., 2nd ed., 1347. 1763, nom. cons.

Epidendropsis Garay & Dunst., in Dunsterville and Garay, Venez. Orchids 

Ill. 6: 39. 1976.

Kalopternix Garay & Dunst., in Dunsterville and Garay, Venez. Orchids Ill. 

6: 40. 1976.

Minicolumna Brieger, in Brieger et al., Orchideen (Schlechter), 3rd ed., 8(29–

32): 495. 1976, nom. illeg.

Neowilliamsia Garay, Orchid Digest 41: 20. 1977.

Takulumena Szlach., in Szlachetko et al., Orchidee (Hamburg) 57: 326. 2006.

Epiphytic, terrestrial or lithophytic herbs, to 3(–7) m tall; 

caespitose; sympodial or rarely monopodial; erect, creeping or 

scandent. Rhizomes short to elongate or absent. Stems usually 

cane-like, simple or branched, rarely thickened into pseudobulbs, 

distichously foliose or with the leaves clustered at the apex. Leaves 

conduplicate, occasionally terete or laterally compressed, slightly 

coriaceous to fleshy, sessile. Inflorescences usually terminal, 

sometimes axillary in leaf sheaths, rarely originating from rhizomes, 1-flowered, racemose or paniculate, sometimes umbellate; 

peduncles naked or sometimes completely or partially covered by 

spathes or imbricate bracts. Flowers resupinate or not, very variable in size and shape (sepals 1–100 mm long), most commonly 

green, white or yellow, but often more brightly colored, usually 

fleshy or subfleshy. Sepals free or rarely somewhat connate, often 

keeled dorsally. Petals free or rarely adnate basally to the column. 

Labellum completely or almost completely fused at the base to 

the apex of the column, free portion simple or lobed, often ornamented with calli or ridges, margins simple or fimbriate, erose 

or denticulate, the portion attached to the column forming a nectary tube; column usually compressed laterally; anther dorsal or 

subterminal, operculate, incumbent, 2- or imperfectly 4-locular; 

pollinaria variable, pollinia 4 or rarely 2 or 6; rostellum horizontal, slit-shaped, and rupturing longitudinally when pollinia are 

removed; viscidium usually well developed; stigma ventral. Ovary 

1-locular. Capsules ellipsoid, pyriform or subspherical, 3-ribbed 

or sometimes winged, smooth or ornamented. [Epidendroideae: 

Epidendreae: Laeliinae.]

A subtropical and tropical American genus of ~1,500 species found in North America (USA), Mexico, Central America, 

the West Indies, and South America (Colombia, Venezuela, Trinidad and Tobago, the Guianas, Ecuador, Peru, Bolivia, Brazil, 

Paraguay, and Argentina); most diverse in the Andes. Approximately 160 species occur in Venezuela.

References. Berg et al. (2009); Dressler (1982, 

1984a, 1984b); Pinheiro and Cozzolino (2013); Pridgeon et al. 

(2005).

In its current circumscription, Epidendrum is a monophyletic, albeit large, genus (Pridgeon et al., 2005). The actual number of species of Epidendrum is difficult to ascertain since until 

recently many species were very broadly circumscribed, particularly in some Andean groups such as the E. cernuum Kunth 

complex. Most species of Epidendrum have dull-colored flowers 

that are fragrant nocturnally, and they appear to be moth pollinated, whereas other species have brightly colored flowers and 

are likely butterfly or hummingbird pollinated.

KEY TO THE SPECIES OF EPIDENDRUM

1a. Stems 1-foliate or rarely 2-foliate, somewhat pseudobulbous-thickened, always composed of 1 internode; the 1 or 2 

leaves apical or subapical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Inflorescences subumbellate, 2–8-flowered, 3–5 cm long; peduncles terete; petals narrowly lanceolate; labellum with 

3 lobes of ± the same shape and size; labellum disk basally with 2 calli; leaf blades elliptic, 3–5× longer than wide

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. jajense

2b. Inflorescences racemes, many-flowered, 15–28 cm long; peduncles and sheaths winged; petals linear; labellum 

cordate-deltoid or shallowly 3-lobed, lateral lobes placed on the basal 1⁄2 of the labellum, perpendicular to the axis 

of the labellum or retrorse; labellum disk with 3 calli or keels; leaf blades linear-elliptic, ~8–15× longer than wide

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. attenuatum

1b. Stems several- to many-leaved, not pseudobulbous-thickened, composed of 3 to many internodes; leaves distichous on the 

stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Inflorescences lateral, subumbellate, several per stem, produced from the middle internodes . . . . . . . E. dendrobii

3b. Inflorescences terminal, racemose or paniculate, 1 per stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Sympodial systems composed of long simple, erect stems that branch only at the base; leaves usually 7 or more 

per individual stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
148 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

 5a. Inflorescences broadly paniculate, 2- or 3-branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 6a. Flowers pink or purple; labellum simple, deeply concave; dorsal sepal 5–8 mm long . . . . . . . E. frutex

 6b. Flowers white or green, labellum 3- or 4-lobed, convex, dorsal sepal ~10 mm long . . . . E. unguiculatum

 5b. Inflorescences simple or, if branched, then branches simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 7a. Leaf blades 3–8(–10) cm wide; dorsal sepal 3.5–6 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

 8a. Leaf blades obovate, 6–8 cm wide, apices obtuse or rounded; leaf sheaths loosely clasping stems; 

labellum shallowly 3-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. lacustre

 8b. Leaf blades elliptic, 3–5(–10) cm wide, apices obtuse or acute; leaf sheaths tightly clasping stems; 

labellum sharply 3-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. leucochilum

 7b. Leaf blades 1–2.5 cm wide; dorsal sepal 0.5–1 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

 9a. Inflorescences nutant, dense racemes; flowers opening ± simultaneously; rachises much longer than 

peduncles; labellar margins entire; calli 2; flowers greenish, yellowish or dull orange, sometimes with 

pink or purplish tinges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

 10a. Plants stout, mostly terrestrial; stems >1 m tall, at least 1 cm in diameter at base (in herbarium specimens); spathes 15–20 cm long; dorsal sepal 11–16 mm long; lateral sepals not or 

very shallowly keeled dorsally; central lobe of labellum oblong, obtuse, truncate or rounded, 

3-keeled longitudinally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. klotzscheanum

 10b. Plants less robust, usually epiphytic or less commonly terrestrial; stems <1 m tall, <0.5 cm in 

diameter at base (in herbarium specimens); spathes 5–9 cm long; dorsal sepal 7–9(15.5) mm 

long; lateral sepals sharply keeled dorsally; central lobe of labellum narrowly triangular, acute, 

not keeled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. alpicola

 9b. Inflorescences erect, dense subumbels; flowers opening successively; occasionally branched and then 

the branches simple and originating from the peduncle below and chronologically posterior to the first 

inflorescence; rachises much shorter than peduncles; labellar margins fimbriate or laciniate . . . . . 11

 11a. Flowers not resupinate; callosity complex, composed of several tubercles and lamellae; flowers 

white or pale yellow (our material), pink or purple . . . . . . . . . . . . . . . . . . . . . . E. secundum

 11b. Flowers resupinate; callosity simple, composed of 3 keels, central keel much longer than lateral 

keels; flowers orange, red, scarlet, yellow or white . . . . . . . . . . . . . . . . . . . . . . E. ibaguense

 4b. Sympodial systems composed of stems that branch above the base, plants often shrub-like; leaves usually 7 or 

fewer per individual stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

 12a. Many-branched, creeping or pendent plants; leaf blades 3–5 cm × 2–6 mm; inflorescences sessile or almost 

sessile, 1-flowered; labellum deeply concave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. repens

 12b. Sparingly branched, usually erect plants, if pendent, then inflorescences multiflowered; leaf blades >4 cm × 

(4–)6–20(–25) mm (if shorter, then inflorescences subumbellate and, if narrower, then at least 5 cm long); inflorescences pedunculate (if peduncles almost absent, then flowers several); labellum concave or convex . . . . 13

 13a. Rachises of inflorescences entirely covered by imbricate floral bracts; ovary pedicellate, partially 

enveloped by floral bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

 14a. Inflorescences much shorter than apical leaves, successively 2–4-flowered; leaf blades 12–25 

mm wide; dorsal sepal 16–18 mm long; petals spathulate, ± as wide as sepals; labellum simple, 

suborbicular-cordate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. cereiflorum

 14b. Inflorescences as long as or longer than apical leaves, simultaneously 6–9-flowered; leaf blades 

3–8 mm wide; dorsal sepal 4–6 mm long; petals filiform, much narrower than sepals; labellum 

deeply 3-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. andinum

 13b. Rachises of inflorescences naked or floral bracts not imbricate; ovary pedicellate, not enveloped by 

floral bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

15a. Sepals ~5 mm long, apically apiculate, not winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

16a. Inflorescences racemose or with 1 or 2 basal branches; floral bracts ~1⁄2 as long as pedicellate ovaries or longer; leaf 

blades 4–8 cm long; labellum deeply concave with entire margins; pedicellate ovaries smooth; petals filiform, much 

narrower than sepals; sepals lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. guaramacalense

16b. Inflorescences subumbellate; floral bracts much shorter than pedicellate ovaries; leaf blades 2–6 cm long; labellum 

flat or concave with margins finely erose; pedicellate ovaries verruculose; petals elliptic, ± as wide as sepals; sepals 

ovate or ovate-elliptic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. caesaris

15b. Sepals ≥5 mm long, terminating or not in a winged dorsal keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17a. Racemes strictly erect, much longer than subtending leaves; flowers not resupinate; labellum concave, ovatetriangular, obtuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
NUMBER 100 • 149

 18a. Flowers pale lavender or purplish; floral bracts 5–7 mm long; inflorescences dense with internodes of rachises 

much shorter than floral pedicels; stems 8–12 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. sp. B

 18b. Flowers greenish-white with some purple on labellum; floral bracts 9–18 mm long; inflorescences less dense 

with internodes of rachises longer than floral pedicels; stems 5–7 mm in diameter . . . . . . . . . . . . . . E. sp. A

17b. Racemes suberect or pendent, shorter to ± as long as subtending leaves; flowers resupinate (or when morphologically not resupinate, then presented in a natural position with the labellum lowermost because of the pendulous 

nature of the inflorescence); labellum convex or flat, subquadrate to broadly oblong . . . . . . . . . . . . . . . . . . . . 19

 19a. Labellum wider than long, lateral lobes larger than middle lobe, orbicular, middle lobe conspicuously 

emarginate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. curtisii

 19b. Labellum ± as wide as long, lateral lobes smaller than middle lobe, trapezoidal or rounded, middle lobe 

apiculate or entire, not emarginate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

 20a. Inflorescences densely 8–20(–40)-flowered, arching or pendent . . . . . . . . . . . . . . . . E. pseudocernuum

 20b. Inflorescences laxly 1–6-flowered, erect to arching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

 21a. Petals more than 3× as long as wide; leaf blades 0.7–2 cm wide . . . . . . . . . . . . . . . . . . . . . . . . 22

 22a. Sepals conspicuously carinate-winged dorsally; inflorescences 1–3(–5)-flowered . . . . . . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. urichianum

 22b. Sepals not carinate-winged dorsally; inflorescences 4–6-flowered . . . . . . . . . . . . . . . E. sp. C

 21b. Petals less than 3× as long as wide; leaf blades 1.7–3.5 cm wide . . . . . . . . . . . . . . E. chioneoides

Epidendrum alpicola Rchb. f., Bonplandia (Hannover) 2: 110. 

1854 (“alpicolum”); Foldats, Fl. Venez. 15(3): 171–172. 

1970 (“alpicolum”); Dunsterville and Garay, Venez. Orchids 

Ill. 6: 116–117. 1976 (“alpicolum”); Romero and Carnevali, 

Orchids Venez., 2nd ed., 1: 205. 2000 (“alpicolum”); Fernández, Orquídeas Nat. Táchira 68. 2003 (“alpicolum”).

Epiphytic or rarely terrestrial herbs; stems 50–70 cm tall. 

Leaf blades 8–15(–20) × 0.9–2.5 cm, apices acute. Inflorescences 

nutant, 10–15 cm long. Flowers resupinate, pale dull orange. 

Dorsal sepal 7–9(–15.5) mm long.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, the species has a disjunct distribution 

with records from the Andes (Mérida, Táchira, and Trujillo) and 

the Venezuelan Guayana (Amazonas; Map 157). In the park, 

found on the north slope along the Boconó–Guaramacal road; 

2,200 m.

♦ Epidendrum andinum Carnevali & G. A. Romero, in Romero 

and Carnevali, Orchids Venez., 2nd ed., 3: 1131. 2000 (description); ibid., 2nd ed., 1: 209. 2000 (illustration); Fernández, Orquídeas Nat. Táchira 68. 2003.

Epidendrum bifarium auct., non Sw.; Dunsterville and Garay, Venez. Orchids 

Ill. 3: 84–85. 1965; Foldats, Fl. Venez. 15(3): 179–181, fig. 442. 1970; 

Ortega et al., BioLlania 5: 47. 1987; Dorr et al., Contr. U.S. Natl. Herb. 

40: 49. 2000 [2001].

Epiphytic herbs, pendent or arching; stems 15–35 cm long. 

Leaf blades narrowly lanceolate, 2–7 cm × 3–8 mm, apices acute. 

Inflorescences pendent, 6–10-flowered. Flowers simultaneous; 

MAP 157. Epidendrum alpicola occurrence in Venezuela. MAP 158. Epidendrum andinum occurrence in Venezuela.
150 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

perianth segments spreading, yellow, translucent yellow or 

yellowish-green with maroon tinges. Dorsal sepal 4–6 mm long.

Found in South America (Colombia and Venezuela). In Venezuela, only known from the Andes (Táchira and Trujillo; Map 

158). Cloud forest along the Boconó–Guaramacal road and on 

the north slope of Guaramacal; 1,800–2,600 m.

This species is well characterized by its elongated inflorescences with a zigzag rachis completely obscured by floral bracts. 

It is closely related to Epidendrum bifarium Sw. from Jamaica, 

but E. andinum has smaller flowers, and the midlobe of the labellum is not 2-lobed.

Epidendrum attenuatum Lindl., Fol. Orchid. 3: Epidendrum 41. 

1853, non Lindl., Fol. Orchid. 3: Epidendrum 69. 1853; 

Dunsterville and Garay, Venez. Orchids Ill. 4: 76–77. 1966; 

Foldats, Fl. Venez. 15(3): 176–179, fig. 441. 1970; Romero 

and Carnevali, Orchids Venez., 2nd ed., 1: 212. 2000; 

Carnevali and Ramírez-Morillo, in Berry et al., Fl. Venez. 

Guayana 7: 334. 2003; Fernández, Orquídeas Nat. Táchira 

70. 2003; Morillo, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 1: 135–136, fig. 27. 2011.

Epiphytic herbs; stems to 30 cm tall; pseudobulbs present. 

Leaf blades linear-elliptic, 4–12 × 0.3–0.7 cm, apices acute. Inflorescences erect or arching, many-flowered, 15–28 cm long. 

Perianth segments widely spreading or subreflexed, green or 

yellow-green. Sepals 6–7.5 mm long.

Found in South America (Venezuela and Ecuador). Fairly 

widespread in Venezuela, occurring in the Andes (Mérida, 

Táchira, and Trujillo), the Cordillera de la Costa (Distrito Federal and Miranda), and the Venezuelan Guayana (Bolívar; Map 

159). Cloud forest along the Boconó–Guaramacal road; 2,000–

3,000 m.

Epidendrum attenuatum is easy to recognize because of its 1-

or 2-leaved pseudobulbs and elongated racemose inflorescences.

Epidendrum caesaris Hágsater & E. Santiago, Icon. Orchid. 

(Mexico) 9(6): t. 915. 2007; Morillo, in Morillo et al., Bot. 

Ecol. Monocot. Páramos Venez. 1: 136–137, fig. 28. 2011.

Epidendrum globiflorum auct., non F. Lehm. & Kraenzl.; Foldats, Fl. Venez. 

15(3): 262–264, fig. 265. 1970; Ortega et al., BioLlania 5: 47. 1987.

Epidendrum restrepoanum auct., non A. D. Hawkes; Dunsterville and Garay, 

Venez. Orchids Ill. 4: 98–99. 1966; Romero and Carnevali, Orchids 

Venez., 2nd ed., 1: 289. 2000; Dorr et al., Contr. U.S. Natl. Herb. 40: 

50. 2000 [2001]; Fernández, Orquídeas Nat. Táchira 89. 2003.

Epiphytic or subterrestrial herbs; stems 20–30 cm tall. Leaf 

blades narrowly lanceolate, 2–6 cm × 4–11 mm, apices emarginate. Inflorescences subumbellate, few-flowered. Flowers subcampanulate, yellow or brownish with pinkish tinges. Dorsal sepal 

5–7 mm long.

Found in South America (Colombia and Venezuela). In Venezuela, restricted to the Andes (Lara, Mérida, Táchira, and Trujillo; Map 160). Cloud forest, subpáramo, and páramo especially 

on the north slope of Guaramacal; 2,200–3,000 m.

This species has been confused with Epidendrum globiflorum F. Lehm. & Kraenzl., which is endemic to Ecuador, and with 

E. restrepoanum A. D. Hawkes, which occurs in Colombia, Venezuela, and Ecuador. Epidendrum caesaris and E. restrepoanum

are sympatric (Hágsater and Santiago, 2007) but not so far as 

we know in Guaramacal. According to Hágsater and Santiago 

(2007), E. caesaris can be distinguished from E. restrepoanum

by flower color. The former has translucent pale pink flowers 

with the base of the sepals and petals yellow (similar to our material), whereas the latter has scarlet red flowers with a yellow 

labellum and column.

MAP 159. Epidendrum attenuatum occurrence in Venezuela. MAP 160. Epidendrum caesaris occurrence in Venezuela.
NUMBER 100 • 151

Epidendrum cereiflorum Garay & Dunst., in Dunsterville and 

Garay, Venez. Orchids Ill. 3: 88. 1965; Foldats, Fl. Venez. 

15(3): 188–190, fig. 445. 1970; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 218. 2000; Fernández, Orquídeas 

Nat. Táchira 72. 2003.

Epiphytic herbs, straggling or creeping; stems to 1.2 m long. 

Leaf blades 5–15 cm × 12–25 mm, apices acute to acuminate. 

Inflorescences 2–4-flowered, 5–7 cm long. Flowers successive, 

green. Dorsal sepal 16–18 mm long.

Endemic to Venezuela, where it is found in the Andes (Lara, 

Portuguesa, Táchira, and Trujillo) and in the Cordillera de la 

Costa (Distrito Federal and Miranda; Map 161). Cloud forest 

along the Boconó–Guaramacal road; ~2,000 m.

Epidendrum chioneoides Carnevali & G. A. Romero, in Romero 

and Carnevali, Orchids Venez., 2nd ed., 3: 1131. 2000 (description); ibid., 2nd ed., 1: 220. 2000 (illustration); Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 

1: 139–140, fig. 30. 2011.

Epidendrum mojandae auct., non Schltr.; Dunsterville and Garay, Venez. Orchids Ill. 1: 120–121. 1959 (“moyandae”); Foldats, Fl. Venez. 15(3): 

318–319, fig. 498. 1970, pro parte, excluding type & fig. 498; Dunsterville and Garay, Orchids Venez., ed. 1, 1: 258. 1979 (“moyandae”).

Terrestrial herbs; stems to 70 cm long. Leaf blades lanceolate, 

4.5–5.5 cm × 17–35 mm, apices blunt to acute. Inflorescences erect 

to arching, few-flowered. Flowers yellow. Sepals 8–11 mm long.

Endemic to Venezuela, where it is found in the Cordillera 

de Mérida (Trujillo) and in the Cordillera de la Costa (Distrito 

Federal; Map 162). Fire-induced páramo below the Páramo 

del Pumar and along the track from El Campamento to Qda. 

Honda; 1,900–2,600 m.

Epidendrum curtisii A. D. Hawkes, Orquídea (Rio de Janeiro) 

18: 170. 1957; Romero and Carnevali, Orchids Venez., 

2nd ed., 1: 232. 2000, nom. nov. Epidendrum lansbergii

Rchb. f., Ned. Kruidk. Arch. 4: 316. 1859, non Regel, 1855.

Epidendrum superpositum Garay, Bot. Mus. Leafl. 18: 203, t. 38. 1958; 

Dunsterville and Garay, Venez. Orchids Ill. 1: 136–137. 1959; Foldats, 

Fl. Venez. 15(3): 415–416, fig. 537. 1970.

Epiphytic herbs; stems to 15–40 cm tall. Leaf blades 2.6–7 

× 0.8–1.8 cm, apices acute. Inflorescences suberect or pendent, 

MAP 161. Epidendrum cereiflorum occurrence in Venezuela. MAP 162. Epidendrum chioneoides occurrence in Venezuela.

MAP 163. Epidendrum curtisii occurrence in Venezuela.
152 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

5–15-flowered. Flowers resupinate, green, suffused on back with 

light maroon. Dorsal sepal 8–10.5 mm long.

Endemic to Venezuela, where it is found in the Cordillera de 

Mérida (Trujillo) and in the Cordillera de la Costa (Aragua, Distrito Federal, and Miranda; Map 163). Collected once between 

Boconó and Batatal; 1,800 m.

Epidendrum dendrobii Rchb. f., Linnaea 22: 841. 1849; Dunsterville and Garay, Venez. Orchids Ill. 5: 82–83. 1972; Foldats, Fl. Venez. 15(3): 226–228, fig. 461. 1970; Romero and 

Carnevali, Orchids Venez., 2nd ed., 1: 234. 2000; Fernández, Orquídeas Nat. Táchira 74. 2003.

Epidendrum pileatum Rchb. f., Bonplandia (Hannover) 3: 68. 1855.

Terrestrial herbs; stems 0.5–2.5 m tall. Leaf blades narrowly 

lanceolate, 9–16 × 0.8–2.7 cm, apices attenuate. Inflorescences 

lateral, subumbellate, few-flowered. Flowers green, several opening simultaneously. Dorsal sepal 7–9 mm long.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). In Venezuela, restricted to the Andes (Mérida, 

Táchira, and Trujillo; Map 164). In the park, found in forest in 

Qda. Jirajara on the south slope of Guaramacal; 1,900–2,900 m.

When encountered, Epidendrum dendrobii makes an attractive display on road cuts and often attains fairly large dimensions 

vegetatively. It is found at much lower elevations (700 m) in Ecuador than it is in Colombia and Venezuela (1,700–2,900 m).

The flowers of this species are spicily fragrant at night, suggesting moth (or bat?) pollination.

Epidendrum frutex Rchb. f., Xenia Orchid. 1: 95, t. 37. 1855; 

Foldats, Fl. Venez. 15(3): 255–257. 1970, pro parte, excluding fig. 471; Dunsterville and Garay, Venez. Orchids Ill. 6: 

124–125. 1976; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 147. 2011.

Epidendrum frigidum auct., non Linden ex Lindl.; Romero and Carnevali, 

Orchids Venez., 2nd ed., 1: 242. 2000.

Terrestrial herbs; stems robust, erect, 1–3 m tall. Leaf 

blades narrowly lanceolate, 10–20 × 0.8–3 cm, apices acute. 

Inflorescences large, broadly paniculate, 2- or 3-branched, pendent. Flowers campanulate, opening in slow succession on the 

branches of the panicle, pale pinkish-purple to dull purple. Dorsal sepal 6–9 mm long.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, this species occurs in the Andes (Táchira 

and Trujillo) and in the Sierra de Perijá (Zulia; Map 165). In 

the park, found in subpáramo and páramo on both slopes of 

Guaramacal; 2,800–3,100 m.

This is a large species commonly found in páramo and subpáramo in the northern Andes. It is easily recognized by its 2- or 

3-pinnate, pendent panicles of pinkish or purplish flowers.

♦ Epidendrum guaramacalense Hágsater, Icon. Orchid. (Mexico) 7(4): t. 740. 2004; Morillo, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 1: 147–148. 2011.

Epidendrum aymardiorum Carnevali & I. Ramírez, nom. nud., in sched.; 

Dorr et al., Contr. U.S. Natl. Herb. 40: 49. 2000 [2001].

Terrestrial herbs; stems to 50 cm tall. Leaf blades lanceolate, 

4–8 cm × 5–8 mm, apices acuminate. Inflorescences racemes with 

1 or 2 basal branches, arched, few-flowered, to 3.7 cm long. Flowers campanulate, light green or yellow. Sepals 4.5–5 mm long.

Endemic to the Andes of Venezuela, where it is known only 

from the Cordillera de Mérida (Portuguesa and Trujillo; Map 

166). Found in subpáramo on the Boconó–Guaramacal road and 

in disturbed páramo below Páramo del Pumar; 1,950–3,000 m.

MAP 165. Epidendrum frutex occurrence in Venezuela.

MAP 164. Epidendrum dendrobii occurrence in Venezuela.
NUMBER 100 • 153

This species belongs in the taxonomically difficult Epidendrum elleanthoides Schltr. complex. Epidendrum guaramacalense can be recognized within this group by stems that branch 

above midheight, relatively large leaf blades, and relatively long 

linear or linear-elliptic floral bracts.

Epidendrum ibaguense Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 

1: 282. 1815 [1816]; ibid. [qu.] 1: 352. 1815 [1816] (“ybaguense”); Dunsterville and Garay; Venez. Orchids Ill. 2: 96, 

97, 128–129. 1961; Foldats, Fl. Venez. 15(3): 277–279. 

1970; Carnevali and G. Romero, Orchids Venez., 2nd ed., 

1: 246. 2000; Carnevali and Ramírez-Morillo, in Berry et 

al., Fl. Venez. Guayana 7: 338, fig. 325. 2003; Fernández, 

Orquídeas Nat. Táchira 80. 2003.

Terrestrial or lithophytic, rarely epiphytic herbs; stems erect 

or pendulous, to 1 m long. Leaf blades oblong-elliptic, 1.5–16 

× 0.6–5.6 cm, apices rounded to obtuse. Inflorescences erect, to 

70 cm long. Flowers resupinate, orange, red, scarlet, yellow or 

white. Sepals 7–22 mm long.

Found in Mexico, Central America, and South America (Colombia, Venezuela, Guyana, French Guiana, Peru, Bolivia, and Brazil). Widespread in Venezuela (Amazonas, Aragua, Bolívar, Falcón, 

Lara, Mérida, Táchira, Trujillo, and Zulia; Map 167). Found below 

the lower boundary of the park on the north slope in Qda. Chandá, 

but probably present within the boundaries of the park; 1,350 m.

Epidendrum jajense Rchb. f., Bonplandia (Hannover) 2: 20. 

1854; Foldats, Fl. Venez. 15(3): 289–290. 1970; Dunsterville and Garay, Venez. Orchids Ill. 6: 130–131. 1976; 

Romero and Carnevali, Orchids Venez., 2nd ed., 1: 250. 

2000; Fernández, Orquídeas Nat. Táchira 82. 2003.

Epidendrum breviracemum C. Schweinf., Bot. Mus. Leafl. 15: 139, t. 41. 

1952.

Epiphytic herbs; stems 5–12 cm tall; pseudobulbs present. 

Leaf blades elliptic, 4.5–8 × ~1.3 cm, apices acute or subacute. 

Inflorescences subumbellate, 2–8-flowered, 3–5 cm long. Flowers with widely spreading segments, yellow-green. Dorsal sepal 

10–15 mm long.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). Venezuelan material is from the Andes (Lara, 

Mérida, Táchira, and Trujillo; Map 168). Cloud forest on the 

north slope of Guaramacal; 2,100 m.

MAP 167. Epidendrum ibaguense occurrence in Venezuela. MAP 168. Epidendrum jajense occurrence in Venezuela.

MAP 166. Epidendrum guaramacalense occurrence in Venezuela.
154 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

This is an interesting species characterized by its apically 

1-leaved, thinly fusiform stems and subumbellate inflorescences. 

Epidendrum jajense resembles some species of Pleurothallis or 

species of Prosthechea Knowles & Westc.

Epidendrum jajense typically turns a distinctive black color 

when material is pressed and dried for the herbarium.

Epidendrum klotzscheanum Rchb. f., Linnaea 22: 838. 1849; 

Dunsterville and Garay, Venez. Orchids Ill. 3: 102–103. 

1965; Foldats, Fl. Venez. 15(3): 292–295, fig. 486. 1970; 

Romero and Carnevali, Orchids Venez., 2nd ed., 1: 251. 

2000; Carnevali and Ramírez-Morillo, in Berry et al., Fl. 

Venez. Guayana 7: 338–339. 2003; Morillo, in Morillo et 

al., Bot. Ecol. Monocot. Páramos Venez. 1: 150–152, fig. 

37, foto 19. 2011.

Terrestrial herbs; stems 1.5–3 m tall. Leaf blades oblong or 

oblong-lanceolate, 9–25 × 1.6–4 cm, apices acute or obtuse. Inflorescences nutant, densely many-flowered. Flowers with widely 

spreading perianth segments, creamy green, often with pink or 

purplish tinges. Dorsal sepal 11–16 mm long.

Found in South America (Colombia, Venezuela, Ecuador, 

and Brazil). In Venezuela, found in the Andes (Lara, Mérida, 

Táchira, and Trujillo), the Cordillera de la Costa (Anzoátegui, 

Aragua, Distrito Federal, and Sucre), and the Venezuelan Guayana (Amazonas; Map 169). On road cuts in cloud forest on the 

south slope of Guaramacal; 1,800–3,000 m.

Epidendrum lacustre Lindl., Fol. Orchid. 3: Epidendrum 50. 

1853; Dunsterville and Garay, Venez. Orchids Ill. 5: 92–93. 

1972; Foldats, Fl. Venez. 15(3): 295–297, fig. 487. 1970; 

Romero and Carnevali, Orchids Venez., 2nd ed., 1: 252. 

2000; Fernández, Orquídeas Nat. Táchira 83. 2003.

Terrestrial or epiphytic herbs; stems erect, to 1.2 m 

tall. Leaf blades obovate, 20–25 × 6–8 cm, apices obtuse or 

rounded. Inflorescences erect, lax, to 40 cm long; spathe subtending inflorescences 10–15 cm long. Flowers white. Sepals 

3.5–6 cm long.

Found in Central America and South America (Colombia, 

Venezuela, Ecuador, and Peru). In Venezuela, confined to the 

Andes (Mérida, Táchira, and Trujillo; Map 170). Found between 

El Campamento below Cerro El Diablo and Maciegal; 1,800 m.

Carnevali et al. in Hokche et al. (2008) stated that this species was endemic to Venezuela, but that is not the case.

Epidendrum leucochilum Link et al., Allg. Gartenzeitung 11: 

145. 1843; Dunsterville and Garay, Venez. Orchids Ill. 1: 

114–115. 1959; Foldats, Fl. Venez. 15(3): 301–304, fig. 

490. 1970; Dunsterville and Garay, Orchids Venez., ed. 1, 

1: 246. 1979; Carnevali and Ramírez-Morillo, in Berry et 

al., Fl. Venez. Guayana 7: 339. 2003; Fernández, Orquídeas 

Nat. Táchira 84. 2003; Llamozas et al., Libro Rojo Fl. 

Venez. 418. 2003.

Epidendrum longiflorum Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 284. 

1815 [1816]; ibid. [qu.] 1: 354. 1815 [1816], non J. Koenig, 1791; 

Romero and Carnevali, Orchids Venez., 2nd ed., 1: 255. 2000; Dorr et 

al., Contr. U.S. Natl. Herb. 40: 50. 2000 [2001].

Epiphytic or terrestrial herbs; stems erect, 60–120 cm tall. 

Leaf blades elliptic, 10–15(–30) × 3–5(–10) cm, apices obtuse 

or acute. Inflorescences erect, lax, 3–7(–12 or more)-flowered. 

Flowers resupinate, simultaneous. Sepals 4.5–5.5 cm long.

Found in South America (Colombia, Venezuela, and Ecuador). In Venezuela, known from the Andes (Barinas, Lara, 

Mérida, Táchira, and Trujillo), the Sierra de Perijá (Zulia), and 

MAP 170. Epidendrum lacustre occurrence in Venezuela.

MAP 169. Epidendrum klotzscheanum occurrence in Venezuela.
NUMBER 100 • 155

the Cordillera de la Costa (Aragua and Distrito Federal; Map 

171). Laguna de Aguas Negras; 1,900 m.

There is a doubtful record of this species occurring in the 

Venezuelan Guayana (Carnevali and Ramírez-Morillo in Berry et 

al., 2003: 339). The record is not mapped here as it is based on 

suspect locality data associated with a living collection cultivated 

in Mexico.

♦ Epidendrum pseudocernuum Carnevali & I. Ramírez, Harvard Pap. Bot. 3: 241, fig. 1. 1998; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 280. 2000; Fernández, 

Orquídeas Nat. Táchira 88. 2003; Morillo, in Morillo et 

al., Bot. Ecol. Monocot. Páramos Venez. 1: 159, fig. 42, 

foto 27. 2011.

Epidendrum cernuum auct., non Kunth; Dunsterville and Garay, Venez. Orchids Ill. 3: 90–91. 1965; Foldats, Fl. Venez. 15(3): 190–192, fig. 446. 

1970; Ortega et al., BioLlania 5: 47. 1987; Dorr et al., Contr. U.S. Natl. 

Herb. 40: 49. 2000 [2001].

Terrestrial or epiphytic herbs; stems 0.5–1 m tall. Leaf 

blades oblong, 4.5–11 × 1–2.5 cm, apices obtuse. Inflorescences 

arching or pendent, 8–20(–40)-flowered. Flowers with widely 

spreading perianth segments, pale green or creamy white. Dorsal 

sepal 12–14 mm long.

Endemic to the Andes of Venezuela (Barinas, Mérida, 

Táchira, and Trujillo; Map 172). Cloud or elfin forest and 

páramo, often on steep road cuts but certainly also as an epiphyte, 

both slopes of Guaramacal and in the Páramo de Guaramacal; 

2,000–3,100 m.

Epidendrum pseudocernuum is one of several closely related taxa in the E. cernuum species complex. Within the complex, our species is characterized by the oblong, truncate central 

lobe of the labellum that has a subapical apiculate tooth and 

relatively small, reniform or transversely ovate, falcate lateral 

lobes placed in a basal position. Epidendrum cernuum s. str. has 

an acuminate, proportionally narrower central lobe of the labellum; it is also a taxon from the western slopes of the Andes in 

Colombia and Ecuador. The two populations currently assigned 

to E. pseudocernuum are somewhat variable in flower size, 

the degree to which the lateral sepals are subapically keeled, 

and subtle details of the shape of the lateral lobes, but for both 

populations the central lobe is consistent in shape and general 

morphology. A thorough revision of the E. cernuum complex is 

warranted.

Epidendrum repens Cogn., Repert. Spec. Nov. Regni Veg. 7: 

122. 1909; Dunsterville and Garay, Venez. Orchids Ill. 4: 

96–97. 1966; Foldats, Fl. Venez. 15(3): 365–366, fig. 515. 

1970; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 

288. 2000; Carnevali and Ramírez-Morillo, in Berry et al., 

Fl. Venez. Guayana 7: 342. 2003.

Small, epiphytic, pendulous or creeping herbs; stems to 60 

cm long. Leaf blades linear-oblong, 3–5 cm × 2–6 mm, apices 

obtuse. Inflorescences 1-flowered, sessile or almost sessile. Flowers solitary in apices of branches, green or cream-colored. Sepals 

8–13 mm long.

Found in Mexico, Central America, the Greater Antilles, 

and South America (Colombia, Venezuela, Guyana, Ecuador, 

Peru, and Bolivia). In Venezuela, known from the Andes (Lara, 

Mérida, Portuguesa, and Táchira), the Cordillera de la Costa 

(Aragua, Distrito Federal, and Yaracuy), and the Venezuelan 

Guayana (Amazonas and Bolívar; Map 173). La Divisoria de la 

Concepción; 1,700 m.

MAP 171. Epidendrum leucochilum occurrence in Venezuela. MAP 172. Epidendrum pseudocernuum occurrence in Venezuela.
156 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Epidendrum secundum Jacq., Enum. Syst. Pl. 29. 1760; Foldats, 

Fl. Venez. 15(3): 391–395, fig. 527. 1970; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 295. 2000; Carnevali and 

Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 7: 342–

343, fig. 326. 2003; Fernández, Orquídeas Nat. Táchira 92. 

2003.

FIGURE 25I

Terrestrial or rarely epiphytic herbs; stems 0.5–2.5 m tall. 

Leaf blades elliptic, 5–9 × 1.5–2.5 cm, apices obtuse, retuse or 

emarginate. Inflorescences erect, densely many- to few-flowered. 

Flowers not resupinate; petals and sepals widely spreading or reflexed, white with a bright yellow labellum or pale yellow. Dorsal sepal 9–11 mm long.

Found in Mexico, Central America, the Lesser Antilles, and 

South America (Colombia, Venezuela, Trinidad and Tobago, 

Guyana, Ecuador, Peru, Bolivia, Brazil, and Paraguay). Widespread in Venezuela (Amazonas, Anzoátegui, Apure, Bolívar, 

Delta Amacuro, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, 

Trujillo, and Yaracuy; Map 174). On road and trail cuts on both 

slopes of Guaramacal; 1,700–2,100(–3,000) m.

Our material belongs to an entity within the Epidendrum 

secundum complex characterized by narrow, reflexed, white 

perianth segments and a relatively narrow labellum with an irregularly erose or paucidentate apex. The labellum in this species is provided with a relatively small callus, and the keels and 

grooves of this callus are shallow and have smooth surfaces. This 

entity is apparently rare in cloud forest of the Cordillera de la 

Costa and Andean portions of Venezuela, usually occurring as 

an epiphyte, but its distribution outside Venezuela as well as its 

identity are difficult to ascertain because of confusion in the taxonomy of the E. secundum complex.

Epidendrum secundum is visited by butterflies in South 

America, which are presumed to be its principal pollinators 

(Pansarin and Amaral, 2008).

We consider Epidendrum secundum sensu Dunsterville and 

Garay (1966: 74, 1976: 40) to be E. anceps Jacq., a distinct species not found in the Venezuelan Andes.

Epidendrum unguiculatum (C. Schweinf.) Garay & Dunst., in 

Dunsterville and Garay, Venez. Orchids Ill. 6: 38. 1976; 

Romero and Carnevali, Orchids Venez., 2nd ed., 1: 314. 

2000; Carnevali and Ramírez-Morillo, in Berry et al., Fl. 

Venez. Guayana 7: 344. 2003; Fernández, Orquídeas Nat. 

Táchira 98. 2003. Epidendrum paniculatum var. unguiculatum C. Schweinf., Bot. Mus. Leafl. 11: 107. 1943.

Epidendrum paniculatum auct., non Ruiz & Pav.; Foldats, Fl. Venez. 15(3): 

332–335, fig. 503. 1970, pro parte.

Epidendrum fastigiatum auct., non Lindl.; Dunsterville and Garay, Venez. 

Orchids Ill. 2: 118–119. 1961.

Epiphytic or terrestrial herbs; stems to 1 m tall. Leaf 

blades elliptic-ovate, 10–14 × 3–4.5 cm, apices acute to acuminate. Inflorescences broadly paniculate, 2- or 3-flowered, 

exceeding the leaves in size. Perianth segments bright green, 

labellum white with callus and disk purple. Dorsal sepal 12–15 

mm long.

Found in South America (Colombia, Venezuela, Suriname, 

French Guiana, Ecuador, Peru, and Brazil). In Venezuela, collections have been made in Amazonas, Apure, Falcón, Mérida, Miranda, Monagas, Táchira, and Trujillo (Map 175). Our records 

MAP 173. Epidendrum repens occurrence in Venezuela. MAP 174. Epidendrum secundum occurrence in Venezuela.
NUMBER 100 • 157

are from Qda. Segovia and Qda. Honda in sector El Santuario; 

1,800–2,350 m.

Epidendrum urichianum Carnevali et al., Orquídea (Mexico 

City), ser. 2, 12: 151, figs. 1992; Romero and Carnevali, 

Orchids Venez., 2nd ed., 1: 315. 2000.

Epiphytic herbs; stems to 30 cm long. Leaf blades narrowly 

oblong-elliptic to elliptic, 3.8–4.7 × 0.8–1.2 cm, apices rounded 

to obtuse. Inflorescences erect, 1–3(–5)-flowered. Flowers pale 

green or yellow, sometimes suffused with pale lavender. Dorsal 

sepal 7–7.5 mm long.

Found in South America (Venezuela and Guyana). In Venezuela, found in the Cordillera de Mérida (Portuguesa) and the 

Venezuelan Guayana (Amazonas and Bolívar; Map 176). Cloud 

forest near La Divisoria de la Concepción; 1,700 m.

Our record is based on an identification by Hágsater (in 

sched.) of a specimen (van der Werff et al. 7593, MO) collected 

in the park. The specimen clearly belongs in the Epidendrum ramosum Jacq. species complex, as does E. urichianum (described 

from the Venezuelan Guayana), but we are not altogether convinced that our material and the few specimens known from the 

Venezuelan Guayana are identical. Resolution of this problem will 

require further study as E. ramosum s.l. is polymorphic and one 

of the most widespread and common of all neotropical orchids.

Epidendrum sp. A

Epidendrum macrostachyum auct., non Lindl.; Ortega et al., BioLlania 5: 

47. 1987; Dorr et al., Contr. U.S. Natl. Herb. 40: 50. 2000 [2001], 

pro parte.

Terrestrial herbs; stems to 50 cm tall. Leaf blades lanceolate 

to narrowly ovate, 12–15 × 1.2–1.9 cm, apices obtuse. Inflorescences erect, many-flowered, to 15 cm long. Flowers campanulate, 

greenish-white with purple on labellum. Sepals 7.5–8.5 mm long.

Evidently endemic to the Cordillera de Mérida of Venezuela (Trujillo; Map 177). Cloud forest below the Páramo de 

Guaramacal; ~1,500–2,600 m.

This undescribed species belongs to the difficult Epidendrum 

macrostachyum Lindl. complex. Its long floral bracts distinguish 

it from other Venezuelan taxa belonging to this complex.

MAP 176. Epidendrum urichianum occurrence in Venezuela. MAP 177. Epidendrum sp. A occurrence in Venezuela.

MAP 175. Epidendrum unguiculatum occurrence in Venezuela.
158 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Epidendrum sp. B

Epidendrum macrostachyum auct., non Lindl.; Dorr et al., Contr. U.S. Natl. 

Herb. 40: 50. 2000 [2001], pro parte.

Epidendrum obovatipetalum auct., non Hágsater & Dodson; Dorr et al., Contr. U.S. Natl. Herb. 40: 50. 2000 [2001].

Terrestrial herbs; stems 60–100 cm tall. Leaf blades lanceolate to narrowly ovate, 8–12 × 0.2–0.3 cm, apices rounded, minutely apiculate. Inflorescences erect, many-flowered, to 18 cm 

long. Flowers pale lavender or purplish. Sepals 8–11 mm long.

Evidently endemic to the Cordillera de Mérida of Venezuela (Trujillo; Map 178). Cloud forest, often on road cuts and in 

other disturbed areas such as El Campamento in Qda. Jirajara; 

2,200–2,900 m.

Epidendrum sp. C

Epidendrum cardioglossum auct., non Rchb. f.; Dunsterville and Garay, 

Venez. Orchids Ill. 4: 78–79. 1966; Foldats, Fl. Venez. 15(3): 185–188. 

1970, pro parte; Ortega et al., BioLlania 5: 47. 1987; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 213. 2000; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 49. 2000 [2001].

Terrestrial or epiphytic herbs; stems to 1 m tall. Leaf blades 

oblong to linear-oblong, 8–11 × 1–2 cm, apices obtuse. Inflorescences erect, 4–6-flowered. Flowers opening widely, pale green. 

Sepals 8–9 mm long.

Evidently endemic to Venezuela, where it has been found 

only in Guaramacal (Trujillo state; Map 179). Páramo or 

subpáramo at the summit of the Boconó–Guaramacal road; 

~3,000–3,100 m.

This species belongs to an Andean complex that includes 

species such as Epidendrum cernuum, E. fruticetorum Schltr., E. 

mojandae Schltr., E. pichinchae Schltr., and E. pseudocernuum. 

The taxa in this complex are characterized by stems branching 

above the base, forming shrub-like, erect or subscandent plants, 

with short, simultaneously flowered, nutant racemes and ± 3-

lobed labella. This complex seems to be composed of several to 

many locally restricted species, and a careful revision is required. 

The type of E. cardioglossum Rchb. f. is from the Cordillera de la 

Costa and has a sharply 3-lobed labellum, whereas our material 

of E. sp. C displays an ovate-cordate, shallowly lobed labellum.

The following species of Epidendrum is excluded from the 

flora of Guaramacal: Epidendrum vareschii Foldats was attributed to the flora of Guaramacal by Ortega et al. (1987: 47), but at 

present it is known only from the type locality, the Páramo de Boconó (i.e., the Páramo de La Cristalina in Dinira National Park).

Fernandezia Ruiz & Pav.

Fernandezia Ruiz & Pav., Fl. Peruv. Prodr. 123. 1794.

Pachyphyllum Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 271. 1815 [1816]; 

ibid. [qu.] 1: 338. 1815 [1816].

Centropetalum Lindl., Sert. Orch. sub. t. 21. 1839.

Nasonia Lindl., in Bentham, Pl. Hartw. 150. 1844.

Orchidotypus Kraenzl., Bot. Jahrb. Syst. 34: 383. 1906.

Raycadenco Dressler, Icon. Pl. Trop., ser. 2, 6: t. 577. 1989.

Epiphytic, lithophytic or subterrestrial herbs; vegetatively 

reduced. Stems monopodial, simple or branching from the base 

in age, elongate, completely covered by sheaths. Leaves distichous, articulate, conduplicate; leaf blades mostly oblong or 

elliptic, margins usually denticulate, usually succulent. Inflorescences lateral racemes emerging from the axils of leaf sheaths, 

shorter to ± as long as leaves; peduncles abbreviated, subterete or 

MAP 178. Epidendrum sp. B occurrence in Venezuela. MAP 179. Epidendrum sp. C occurrence in Venezuela.
NUMBER 100 • 159

angled; rachises densely or laxly flowered, occasionally abbreviated, and the inflorescences becoming subumbellate; floral bracts 

short. Flowers resupinate, inconspicuous, white or greenish (our 

species), yellow with reddish-brown maculations or bright red, 

red-orange or yellow; perianth segments campanulate or with the 

apical 1⁄2 spreading, margins often ciliate. Sepals free or connate 

in the lower 1⁄2, equal, mostly elliptic or triangular elliptic. Petals 

similar to the sepals, often more obtuse. Labellum free to base, 

enclosed in the sepaline cup, elliptic or oblong, basally clawed or 

sessile, usually pandurate or constricted about midlength, in lateral profile sigmoid, the basal 1⁄2 concave, the apical 1⁄2 recurved, 

disk bicallose or bilamellate; column broadly petaloid, hoodlike, 

easily flattened into a lamina, basally often produced into a small 

foot; anther deeply seated under the hoodlike clinandrium, operculate, 1-locular; pollinia 2, waxy, on narrow, spathulate stipes 

that in turn are united to small, disciform viscidia; stigmatic surface narrow, at the base of the ventral face of the column. Ovary 

cylindrical to trigonous, pedicellate, glabrous. Capsules often 

3-winged. [Epidendroideae: Cymbidieae: Oncidiinae.]

A neotropical genus of ~50 species found in Central America 

and South America (Colombia, Venezuela, Ecuador, Peru, and 

Bolivia). Four species occur in Venezuela, usually above 2,500 m 

elevation in cloud or elfin forest at the limit of páramo.

References. Chase and Whitten (2011); Christenson (2008); Neubig et al. (2012); Pridgeon et al. (2009); Senghas 

(1974, 1995a).

KEY TO THE SPECIES OF FERNANDEZIA

1a. Plants small, >5 cm long; leaf blades 8–12(–15) mm long, leathery; flowers 2.5–3 mm long, perianth segments free . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. crystallina

1b. Plants minute, <5 cm long; leaf blades 3–6 mm long, fleshy-coriaceous; flowers 1.8–2 mm long, basal 1⁄2 of perianth 

segments connate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. schultesii

Fernandezia crystallina (Lindl.) M. W. Chase, in Chase and 

Whitten, Phytotaxa 20: 29. 2011. Pachyphyllum crystallinum Lindl., Orchid. Linden. 18. 1846; Dunsterville and 

Garay, Venez. Orchids Ill. 3: 234–235. 1965; Foldats, Fl. 

Venez. 15(5): 408–409. 1970; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 718. 2000; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 52. 2000 [2001]; Fernández, Orquídeas 

Nat. Táchira 169. 2003; Morillo, in Morillo et al., Bot. 

Ecol. Monocot. Páramos Venez. 1: 240. 2011.

Pachyphyllum pastii Rchb. f., Bonplandia (Hannover) 3: 239. 1855; Foldats, 

Fl. Venez. 15(5): 412–414, fig. 903. 1970; Ortega et al., BioLlania 5: 48. 

1987. Fernandezia pastii (Rchb. f.) M. W. Chase, in Chase and Whitten, 

Phytotaxa 20: 30. 2011.

Epidendrum frutex auct., non Rchb. f.; Ortega et al., BioLlania 5: 47. 1987.

Epiphytic herbs; stems 5–18(–30) cm long. Leaf blades 

8–12(–15) mm long. Inflorescences 2- or 3(5)-flowered. Sepals 

pale-green or yellowish; dorsal sepal ~2 mm long. Petals white; 

labellum white and green toward base.

Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, Ecuador, and Peru). In 

Venezuela, known from the Andes (Mérida, Táchira, and Trujillo) and the Cordillera de la Costa (Distrito Federal, Miranda, 

and Sucre; Map 180). Found in páramo in the park, including 

the Fila de Los Recostaderos; 2,100–3,100 m.

Fernandezia schultesii (L. O. Williams) Carnevali & Dorr, comb. 

nov. Pachyphyllum schultesii L. O. Williams, Caldasia 1(3): 

15, t. p. 16. 1941; Dunsterville and Garay, Venez. Orchids Ill. 

6: 310–311. 1976; Romero and Carnevali, Orchids Venez., 

2nd ed., 3: 720. 2000; Fernández, Orquídeas Nat. Táchira 

170. 2003; Morillo, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 1: 241, fig. 89. 2011. Orchidotypus schultesii (L. O. Williams) Senghas, Orchideen I/B, 31: 1923. 1995.

Pachyphyllum hispidulum auct., non (Rchb. f.) Garay & Dunst.; Foldats, Fl. 

Venez. 15(5): 409–410. 1970, pro parte.

Epiphytic herbs; stems 1–4 cm long. Leaf blades 3–6 mm 

long. Inflorescences 1(2)-flowered. Sepals, petals, and labellum 

fused over 1⁄2 their length into a single tube, white grading to pale 

green at the base. Dorsal sepal ~2 mm long.

Found in South America (Colombia and Venezuela). In Venezuela, found only in the Andes (Táchira and Trujillo; Map 181). 

Forest on the south slope of Guaramacal; 2,000–2,750 m.

We have two collections from the park of this small and 

easily overlooked species; one of the two was found in a mixture 

MAP 180. Fernandezia crystallina occurrence in Venezuela.
160 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

with Trichosalpinx pusilla (Kunth) Luer, one of several small, 

lepanthiform orchids found in Guaramacal.

Gomphichis Lindl.

Gomphichis Lindl., Gen. Sp. Orchid. Pl. 446. 1840.

Terrestrial, subterrestrial, or, more rarely, subepiphytic herbs, 

erect. Roots fleshy, relatively long, fascicled, pubescent. Stems simple, terete, densely leafy basally, loosely leafy apically. Leaves not 

articulate, convolute, herbaceous or subplicate, basal leaves larger, 

often basally attenuate into a pseudopetiole and then leaf base 

sheathing; leaf blades narrowly elliptic to rarely obovate. Inflorescences terminal, erect, loose to densely many-flowered spikes, 

glabrous to variously pubescent; peduncles much longer than rachises, wrapped in several tubular or foliar sheaths; floral bracts 

conspicuous, membranous. Flowers not resupinate, patent or nutant, campanulate; perianth segments free, white, green or yellow 

(labellum white), variously pubescent or ciliate. Sepals subequal, 

erect or patent in the apical 1⁄2, subfleshy. Petals linear to obovate, 

membranous. Labellum more fleshy than other perianth segments, 

free from the column, commonly rhombic-ovate or suborbicular, 

entire or 3-lobed, usually pubescent, disk thickened-fleshy; column clavate, sigmoid, usually pubescent or papillose, without 

wings or foot, apically obtuse; rostellum erect, broad, clinandrium 

well developed, membranous; anther dorsal, erect, subequal to the 

rostellum; pollinia 4, smooth, without caudicles; viscidium terminal; stigma entire. [Orchidoideae: Cranichideae: Cranichidinae.]

Approximately 20 species found in Central America (Costa 

Rica and Panama) and South America (Colombia, Venezuela, 

Guyana, Ecuador, Peru, Bolivia, and Brazil). The center of species diversity for the genus is in the northern Andes of Colombia 

and Venezuela; ~10 species are found in Venezuela.

References. Alvarez (2005); Álvarez-Molina and 

Cameron (2009); Pridgeon et al. (2003); Salazar Chávez et al. 

(2009); Senghas (1994d).

Species of Gomphichis are typically shade tolerant.

Specimens of Gomphichis prepared for the herbarium are 

usually black or dark brown when dry.

There is molecular support for the monophyly of Gomphichis (Álvarez-Molina and Cameron, 2009; Salazar Chávez et al., 

2009). The genus is closely related to several other genera of 

terrestrial orchids found in Central America and the Andes and 

Guayana Highlands of South America, including Aa.

KEY TO THE SPECIES OF GOMPHICHIS

1a. Leaf blades 15–25 cm long, elliptic or oblanceolate; bracts ovate, glabrous, sparsely pubescent at base; petal apex acute, 

glandular-ciliate apically with simple hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. adnata

1b. Leaf blades 20–35(–50) cm long, oblong-lanceolate; bracts lanceolate, pilose; petal apex acute or obtuse, glandular-ciliate 

apically with simple and branched retrorse hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. altissima

Gomphichis adnata (Ridl.) Schltr., Repert. Spec. Nov. Regni 

Veg. Beih. 6: 51. 1919; Morillo, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 1: 181–182, fig. 55B. 2011. 

Stenoptera adnata Ridl., in Thurn, Timehri 5: 205. 1886.

Gomphichis gracilis Schltr., Repert. Spec. Nov. Regni Veg. Beih. 6: 29. 

1919.

Gomphichis steyermarkii Foldats, Acta Bot. Venez. 3: 331, fig. 8A. 1968.

Gomphichis viscosa auct., non (Rchb. f.) Schltr.; Dunsterville and Garay, 

Venez. Orchids Ill. 2: 162–163. 1961; Foldats, Fl. Venez. 15(1): 393–

396, fig. 147. 1969.

Gomphichis costaricensis auct., non (Schltr.) Ames et al.; Romero and 

Carnevali, Orchids Venez., 2nd ed., 1: 347. 2000; Dorr et al., 

Contr. U.S. Natl. Herb. 41: 50. 2000 [2001], pro parte; Carnevali 

and Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 7: 369, fig. 

345. 2003, pro parte.

Terrestrial or rarely epiphytic herbs, to 90(–120) cm tall 

(in flower). Leaf blades elliptic or oblanceolate, 15–25 cm long. 

Flowers white, whitish-green or greenish-yellow. Dorsal sepal 

(4.5–)5–6(–8) mm long. Labellum 5–7 mm long.

Found in South America (Colombia and Venezuela). Widely 

distributed in the mountains of Venezuela (Anzoátegui, Barinas, 

Bolívar, Distrito Federal, Lara, Mérida, Sucre, Táchira, Trujillo, 

and Zulia; Map 182). In the park, collected in subpáramo in 

Qda. Jirajara, in the Páramo de Guaramacal below the television 

MAP 181. Fernandezia schultesii occurrence in Venezuela.
NUMBER 100 • 161

antennas, and near the summit of the Fila de Agua Fría; (2,200–)

2,800–3,000 m.

Earlier, Dorr et al. (2000) confused this species with Gomphichis costaricensis (Schltr.) Ames et al., which Alvarez (2005) 

indicates has been collected in Venezuela only in Henri Pittier 

National Park (Aragua) and on the Chimantá Massif (Bolívar).

Gomphichis altissima Renz, Candollea 11: 253, figs. 5, 7A, a–g. 

1948; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 183, fig. 55A. 2011.

Gomphichis cladotricha Renz, Candollea 11: 256, figs. 5, 7B, h–n. 1948.

Gomphichis costaricensis auct., non (Schltr.) Ames et al.; Dorr et al., Contr. 

U.S. Natl. Herb. 41: 50. 2000 [2001], pro parte.

Terrestrial herbs, to 135(–180) cm tall (in flower). Leaf blades 

oblong-lanceolate, 20–35(–50) cm long. Flowers white or greenish. Dorsal sepal 6–7(–8) mm long. Labellum 6.7–8.5(–10) mm 

long.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, found in the Andes (Mérida, Portuguesa, Táchira, and Trujillo) and in the Sierra de Perijá (Zulia; 

Map 183). In the park, found on both slopes of Guaramacal; 

1,600–2,300(–2,750) m.

Gomphichis altissima is more robust (taller and with larger 

leaves) than G. adnata and generally is found at lower elevations.

The following species of Gomphichis is excluded from the 

flora of Guaramacal: Gomphichis traceyae Rolfe has been collected near Boconó (Trujillo state) at an unspecified locality or 

localities (Renz 7561, 7566), and it is possible that it will be 

found in Guaramacal. It can be distinguished from our species 

by its linear-oblong to elliptic-lanceolate leaf blades; subdense inflorescence; wider than long labellum with rotund lateral lobes; 

broadly elliptic, 3-nerved, always glabrous petals that show 

densely glandular retrorse cilia; and strongly sigmoid column.

Habenaria Willd.

Habenaria Willd., Sp. Pl. 4(1): 5, 44. 1805.

Terrestrial (our species) or semiaquatic herbs, rarely subepiphytic; erect, often ± succulent. Tubers (rhizoids) 1–3, subterranean, aerial portion usually caducous in the dry season 

and plants persisting via basal tubers. Stems terete, completely 

or nearly completely covered by sheaths. Leaves not articulate, 

prefoliation convolute, usually distichous or spirally arranged 

along the stem, in some species rosulate basally or less commonly above the base, occasionally leaves absent or reduced and 

sheath-like; leaf sheaths infundibuliform (funnel shaped), usually 

appressed to the stem, rarely surrounding the stem more loosely, 

usually shorter than the leaf blade; leaf blades usually lanceolate 

to linear-elliptic, sometimes setaceous or suborbicular, membranous to thinly fleshy. Inflorescences terminal racemes or erect, 

lax or dense terminal spikes, usually with few to many flowers opening in succession or ± simultaneously, rarely 1-flowered; 

floral bracts usually ± conspicuous, sometimes spathaceous. 

Flowers resupinate, inconspicuous to relatively large and showy, 

usually green, white or yellowish opaque (or combinations of 

these colors), less commonly (not our species) orange, bright yellow, pink or purple, usually fragrant at night; perianth segments 

free, spreading or dorsal sepal and petals (or part thereof) in the 

form of a cap on the column, fleshy or subfleshy. Sepals subequal 

MAP 182. Gomphichis adnata occurrence in Venezuela. MAP 183. Gomphichis altissima occurrence in Venezuela.
162 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

or lateral ones larger than the dorsal one, usually wider than the 

petals; petals usually 2-lobed, less commonly simple, very rarely 

3-lobed. Labellum usually 3-lobed, less commonly simple basally 

with a conspicuous spur, rarely (usually in teratological cases) 

spur absent, lateral lobes usually as wide or much narrower than 

the central lobe (linear or filiform), rarely wider; column short 

and wide; anther erect, firmly attached to the column, 2-locular, 

each locule projecting over an elongate canal; pollinia 2, sectile, 

with 2 basal, interlocular caudicles; stigmas convex, 2-lobed, the 

lobes often pedunculate. Ovary often conspicuously pedicellate. 

Capsules ellipsoid. [Orchidoideae: Orchideae: Orchidinae.]

A cosmopolitan genus of 500–600 species that is most 

diverse in the tropics, especially in Brazil. In the Americas, 

Habenaria has been collected in North America (USA), Mexico, 

Central America, the West Indies, and South America (Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina, and Uruguay). Twenty-five species are found in 

Venezuela, most of them in the Venezuelan Guayana.

References. Batista et al. (2011, 2013); Kränzlin 

(1893); Pridgeon et al. (2001a); Renz (1992).

The neotropical species of Habenaria form a monophyletic 

group (Batista et al., 2013), and this neotropical clade together 

with certain African species forms a highly supported core Habenaria clade, which includes the Old World type of the genus. It appears that the neotropical species originated in Africa and that the 

radiation in the Americas is relatively recent (Batista et al., 2013).

KEY TO THE SPECIES OF HABENARIA

1a. Leaves linear-oblong to linear-elliptic oblong; stems entirely dark green; flowers mainly green in dense racemes; anterior 

lobe of petal conspicuously shorter than the posterior lobe; lateral lobes of labellum conspicuously shorter than the central lobe; nectary <15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. gollmeri

1b. Leaves elliptic; stems light green with darker green nerves and purple dots or lines; flowers white in loose racemes; anterior lobe of petal as long as or exceeding in length the posterior lobe; lateral lobes of labellum as long as or exceeding in 

length the central lobe; nectary >15 mm long (usually >20 mm long) . . . . . . . . . . . . . . . . . . . . . . . . . . . H. monorrhiza

Habenaria gollmeri Schltr., Repert. Spec. Nov. Regni Veg. Beih. 

6: 27. 1919; Foldats, Fl. Venez. 15(1): 64–66, fig. 17. 1969; 

Fernández, Orquídeas Nat. Táchira 104. 2003; Morillo, 

in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 

189–190, fig. 57. 2011.

FIGURE 25J

Habenaria parviflora auct., non Lindl.; Dunsterville and Garay, Venez. Orchids Ill. 6: 160–161. 1976; Dorr et al., Contr. U.S. Natl. Herb. 40: 50. 

2000 [2001], pro parte.

Terrestrial herbs, to 30 cm tall. Leaf blades linearlanceolate, 6–12 × 8–15 cm. Inflorescences dense racemes. 

Flowers green with a purple spot on the labellum. Dorsal sepal 

2.5–5 × 1.5–4 mm.

Found in South America (Colombia, Venezuela, and Ecuador). In Venezuela, found in the Andes (Mérida, Táchira, and 

Trujillo), the Cordillera de la Costa (Distrito Federal and Miranda), and the Venezuelan Guayana (Amazonas; Map 184). In 

the park, encountered on both slopes of Guaramacal; 1,800–

2,100(–2,600) m.

Habenaria monorrhiza (Sw.) Rchb. f., Ber. Deutsch. Bot. Ges. 3: 

274. 1885; Foldats, Fl. Venez. 15(1): 74–76, fig. 21. 1969; 

Dunsterville and Garay, Venez. Orchids Ill. 2: 170–171. 

1961; Foldats et al., in Berry et al., Fl. Venez. Guayana 7: 

380. 2003; Fernández, Orquídeas Nat. Táchira 105. 2003. 

Orchis monorrhiza Sw., Prodr. 118. 1788.

Habenaria speciosa Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 44. 1835 [1836].

Terrestrial herbs, to 1 m tall. Leaf blades lanceolate to ovate 

or elliptic-oblong, 4–15 × 1.5–4 cm. Inflorescences loose racemes. Flowers white. Dorsal sepal 5–14 × 4–7 mm.

Found in Mexico, Central America, the West Indies, and South 

America (Colombia, Venezuela, Trinidad and Tobago, Ecuador, 

Peru, Bolivia, and Brazil). Widely distributed in Venezuela (Aragua, Barinas, Bolívar, Carabobo, Cojedes, Distrito Federal, Falcón, 

Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, 

Sucre, Táchira, Trujillo, and Zulia; Map 185). Our record is from 

the south slope of Guaramacal near Qda. Honda; 1,950–2,100 m.

This is one of the most common, widely distributed, and 

variable species in the genus. It is possible that several species are 

included in the wide species concept adopted here and in other 

floristic treatments.

MAP 184. Habenaria gollmeri occurrence in Venezuela.
NUMBER 100 • 163

Jacquiniella Schltr.

Jacquiniella Schltr., Repert. Spec. Nov. Regni Veg. Beih. 7: 123. 1920.

Dressleriella Brieger, Orchideen (Schlechter), 3rd ed., 9(33–36): 555. 1977, 

nom. nud. 

Briegeria Senghas, Orchidee (Hamburg) 31: 29. 1980.

Acrorchis Dressler, Orquidía (Mexico City), n.s., 12: 14. 1990.

Epiphytic or lithophytic, rarely subterrestrial, herbs; small 

to medium size; erect to pendulous, caespitose. Rhizomes almost 

absent, thin. Stems terete or somewhat compressed laterally, with 

leaves almost to the base and completely or almost completely 

enveloped by leaf sheaths, straight or ± zigzag in orientation. 

Leaves distichous, articulate, prefoliation duplicative (conduplicate), subterete or laterally compressed in cross section; sheaths 

often rugulose or striate. Inflorescences terminal, solitary or 

fascicled, pedunculate or not, when pedunculate, peduncles flattened and with 1 or 2 bract-like sheaths. Flowers resupinate, inconspicuous, minute to small, usually opening successively, often 

autogamous to cleistogamous; floral bracts relatively large to inconspicuous, acute to obliquely subtruncate, apiculate; pedicels 

short; perianth segments widely spreading to more commonly 

subparallel to the column, usually green but in some species red 

or brown, fleshy. Sepals subequal, free or shortly fused at the 

base, concave. Petals subsimilar to the sepals, sometimes somewhat shorter and narrower. Labellum fused to the base of the 

column or almost completely free, sessile or shortly unguiculate, 

very fleshy, sometimes geniculate and contracted beneath the 

base, entire or 3-lobed apically, concave, usually elliptic or subrhombic; column relatively short, subterete, somewhat dilated 

apically, without a foot; anther terminal or dorsal, operculate, 

incumbent, 2- or 4-locular; clinandrium small, sometimes inflated or elongate, margins entire, erose or denticulate; pollinia 

4, waxy, compressed laterally, attached to a small viscidium; 

rostellum horizontal, slot-shaped, rupturing longitudinally when 

the pollinia are removed; stigma ventral. Ovary often subglobose or ellipsoid, with a cuniculus (i.e., a tubular nectary hidden 

within the floral tube above the ovary). Capsules ovoid, globose 

to ellipsoid. [Epidendroideae: Epidendreae: Laeliinae.]

A neotropical genus of 8 or 9 species found in Mexico, Central 

America, the West Indies, and South America (Colombia, Venezuela, 

Trinidad and Tobago, the Guianas, Ecuador, Peru, and Brazil), with 

a center of diversity in Costa Rica and Panama, where all but 2 of the 

species are found. Three species occur in Venezuela.

References. Berg et al. (2009); Dressler (1982, 

1984a, 1984b); Pridgeon et al. (2005).

Jacquiniella appears to be monophyletic and is closely related 

to Scaphyglottis Poepp. & Endl. (Berg et al., 2009). We fail to see 

the rationale for maintaining the monotypic Acrorchis Dressler 

distinct from Jacquiniella and consider it to be a synonym.

Jacquiniella teretifolia (Sw.) Britton & P. Wilson, Sci. Surv. 

Porto Rico & Virgin Islands 6: 340. 1926; Romero and 

Carnevali, Orchids Venez., 2nd ed., 2: 387. 2000; Carnevali 

and Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 

7: 398, fig. 363. 2003; Fernández, Orquídeas Nat. Táchira 

108. 2003. Epidendrum teretifolium Sw., Prodr. 121. 

1788; Dunsterville and Garay, Venez. Orchids Ill. 1: 138–

139. 1959; Foldats, Fl. Venez. 15(3): 422–425, fig. 550. 

1970. Dressleriella teretifolium (Sw.) Brieger, Orchideen 

(Schlechter), 3rd ed., 9(33–36): 555. 1977, nom. illeg. Briegeria teretifolia (Sw.) Senghas, Orchidee (Hamburg) 31: 30. 

1980.

Epiphytic or more rarely lithophytic herbs; erect or ascending. Leaf blades ± terete, 1.5–5 cm × 1.5–2.5 mm. Inflorescences 

MAP 186. Jacquiniella teretifolia occurrence in Venezuela.

MAP 185. Habenaria monorrhiza occurrence in Venezuela.
164 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

1-flowered, long-pedunculate. Flowers greenish yellow. Sepals 

8–12.5 mm long.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, the Guianas, and Brazil). 

Widespread in Venezuela (Amazonas, Aragua, Barinas, Bolívar, 

Carabobo, Distrito Federal, Lara, Miranda, Monagas, Portuguesa, Sucre, Táchira, and Zulia; Map 186). La Divisoria de la 

Concepción; 1,300–1,500 m.

Lepanthes Sw.

Lepanthes Sw., Nova Acta Regiae Soc. Sci. Upsal., ser. 2, 6: 85. 1799.

Epiphytic, lithophytic or subterrestrial herbs; caespitose; 

creeping, erect or pendulous. Rhizomes very short to elongate. 

Stems sometimes prolific, 1-leaved apically, densely or laxly 

covered by a series of tubular, ribbed, ± imbricate sheaths with 

oblique, dilated ostia or mouths (i.e., lepanthiform sheaths), 

margins of ostia visibly or microscopically ciliate or scabrous, 

occasionally glabrous. Leaves conduplicate; leaf blades mostly 

flat, sometimes concave or convex, linear-elliptic to orbicular, 

membranous or rarely subfleshy, petiolate. Inflorescences 1 to 

several, originating from an annulus near stem apex, few- to 

many-flowered racemes; rachises straight to fractiflex; floral 

bracts small to conspicuous. Flowers not resupinate, usually 

small, several, opening singly or more commonly successively, 

often lasting 1 day, commonly brightly colored. Sepals glabrous 

or ciliate, usually dorsally carinate; dorsal sepal entirely free to 

variously connate with basal portion of synsepal; lateral sepals 

usually fused into a synsepal, rarely almost free, membranous. 

Petals generally wider than long, very rarely longer than wide 

and then linear or very narrowly triangular, commonly differentiated into distinct upper and lower lobes, upper lobe pointing 

toward dorsal sepal, lower lobe pointing toward lateral sepals, 

morphological apex of petals flat, convex, apiculate concave or 

emarginate. Labellum usually 3-lobed, very rarely entire; central 

lobe modified into a variously shaped and sculptured appendix; 

lateral lobes often peltate or semilunate and embracing the column; base of labellum connate to the under surface of column, 

often near its base; column footless, usually subcylindrical, very 

short to relatively elongate; anther operculate, apical, subapical 

or ventral; pollinia 2, attached to a small viscidium; stigma apical, subapical or ventral. Ovary terete or 3-angled, pedicellate. 

Capsules ellipsoid to obovoid. [Epidendroideae: Epidendreae: 

Pleurothallidinae.]

A neotropical genus found in Mexico, Central America, the 

West Indies, and South America (Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, and Brazil). There are 600–800 species; ~50 are found in Venezuela.

References. Luer (1986a, 1986b); Pridgeon and 

Chase (2001); Pridgeon et al. (2001b, 2005).

Although there appears to be support for the monophyly 

of Lepanthes, the support is based on molecular sequences from 

only four species (Pridgeon and Chase, 2001; Pridgeon et al., 

2001b). Morphologically, lepanthiform sheaths unite Lepanthes, 

Lepanthopsis (Cogn.) Ames, and Trichosalpinx Luer, and floral 

characters (see Subkey V) can be used to separate these three 

closely related genera.

Lepanthes is composed of many narrow endemics differing 

from one other in structural details of the petal and labellum 

shape. The genus is most diverse in the northern Andes, where 

the majority of the species occur in cloud or otherwise humid 

forest at intermediate or high elevations. Few species of Lepanthes grow at low elevations or in dry forest.

In Costa Rica, Blanco and Barboza (2005) observed pseudocopulatory pollination in Lepanthes glicensteinii Luer by a fungus gnat (Diptera: Sciaridae). They postulate that pollination by 

sexual deception is prevalent in the genus because most species 

of Lepanthes have a similar flower structure.

KEY TO THE SPECIES OF LEPANTHES

1a. Inflorescences densely flowered; floral bracts equal to or longer than rachis internodes and the ovary (at least the portion 

that remains on the rachis after flower dehiscence) much longer than rachis internodes . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Leaf blades elliptic or ovate-elliptic, apices obtuse or acute; sepals ~3 mm long; petals asymmetrical, anterior and 

posterior lobes different . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. wageneri

2b. Leaf blades suborbicular, apices rounded; sepals 3.5–4 mm long; petals symmetrical, anterior and posterior lobes 

similar (each petal resembling a ± regular diamond shape) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. sp. A

1b. Inflorescences laxly flowered; floral bracts and the ovary shorter than rachis internodes . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Peduncles exceeding the leaves in length; flowers remote, successive . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. sp. B

3b. Peduncles shorter than or subequal to the leaves in length; flowers ± clustered, ± simultaneous . . . . . . . . . . . . . . 4

 4a. Apex of both lobes of the petals long-acuminate; leaf blades narrowly lanceolate, at least 3.5× longer than wide, 

apices long-acuminate-caudate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. glochidea

 4b. Apex of the petals rounded or obtuse; leaf blades elliptic, 2 or 3× longer than wide, apices acute to obtuse . . . . 5

 5a. Dorsal sepal ~2× longer than wide, <3.5 mm long; appendix of labellum several times longer than wide; attachment point of labellum to column well above the base of the lateral lobes; clinandrium with a tooth in 

the center of the emargination; petals and labellum finely papillose; flowers resupinate . . . . . . . . L. scolex

 5b. Dorsal sepal at least 3× longer than wide, >4 mm long; appendix of labellum broadly ovate or suborbicular, ± 

as long as wide; attachment point of labellum to column at the base of the lateral lobes; clinandrium without 

a tooth in the emargination; petals and labellum ciliolate; flowers not resupinate . . . . . . . . . . L. vareschii
NUMBER 100 • 165

♦ Lepanthes glochidea Luer, Phytologia 59: 446. 1986; Romero 

and Carnevali, Orchids Venez., 2nd ed., 2: 411. 2000.

Lepanthes millei auct., non Schltr.; Dunsterville and Garay, Venez. Orchids 

Ill. 4: 124–145. 1966; Foldats, Fl. Venez. 15(2): 484–485, fig. 372. 

1970; Ortega et al., BioLlania 5: 48. 1987; Morillo, in Morillo et al., 

Bot. Ecol. Monocot. Páramos Venez. 1: 198, fig. 632. 2011.

Epiphytic herbs, 7.5–11 cm tall. Leaf blades narrowly lanceolate, 2–4 × 0.5–0.8 cm. Flowers not resupinate. Sepals yellowish with reddish tinges; dorsal sepal 3.5–4.1 mm long, narrowly 

ovate, apices acuminate. Petals yellow-orange with brownish 

tinges. Labellum brownish or reddish-purple.

Endemic to the Andes of Venezuela (Táchira and Trujillo; 

Map 187). Cloud forest along the Boconó–Guaramacal road; 

1,800–2,750 m.

Lepanthes millei Schltr. is an Ecuadorian species with 

narrowly obovate leaves and narrower petals with obtuse or 

rounded lobes.

♦ Lepanthes scolex Luer, Phytologia 59: 450. 1986; Romero and 

Carnevali, Orchids Venez., 2nd ed., 2: 431. 2000; Fernández, Orquídeas Nat. Táchira 122. 2003.

Lepanthes acuminata auct., non Schltr.; Dunsterville and Garay, Venez. Orchids Ill. 4: 118–119. 1966.

Lepanthes lindleyana auct., non Oerst. & Rchb. f.; Foldats, Fl. Venez. 15(2): 

480–482, fig. 370. 1970, pro parte; Ortega et al., BioLlania 5: 47. 1987 

(as to Dunsterville 724).

Lepanthes stenophylla auct., non Schltr.; Dunsterville and Garay, Venez. Orchids Ill. 2: 184–185. 1961; Foldats, Fl. Venez. 15(2): 503–504, fig. 

382. 1970.

Epiphytic herbs, 3–5 cm tall. Leaf blades elliptic or narrowly elliptic, 1–2 × 0.3–0.9 cm. Flowers resupinate. Dorsal 

sepal 2.8–3.1 mm long, ovate, apices shortly acuminate.

Endemic to the Andes of Venezuela (Táchira and Trujillo; 

Map 188). Cloud forest along the Boconó–Guaramacal road; 

~2,000 m.

This species belongs to the widespread and taxonomically 

difficult Lepanthes lindleyana Oerst. & Rchb. f. complex, but 

the small, elongate, hirsute appendix of the labellum readily sets 

this species apart from its closest relatives. In the park, L. lindleyana could be confused with L. vareschii Garay as sterile material of these two species is indistinguishable. However, the floral 

characters given in the key permit ready identification.

Lepanthes vareschii Garay, Canad. J. Bot. 34: 252, fig. 11A. 

1956; Dunsterville and Garay, Venez. Orchids Ill. 3: 156–

157. 1965; Foldats, Fl. Venez. 15(2): 516–517, 387A. 1970; 

Romero and Carnevali, Orchids Venez., 2nd ed., 2: 439. 

2000; Fernández, Orquídeas Nat. Táchira 120. 2003.

Epiphytic herbs, 3–9 cm tall. Leaf blades elliptic or narrowly elliptic, 1–4 cm × 6–9 mm. Flowers not resupinate, 

yellowish. Dorsal sepal 4–4.5 mm long, ovate, apices shortly 

acuminate.

Endemic to the Andes of Venezuela (Mérida, Táchira, and 

Trujillo; Map 189). Cloud forest along the Boconó–Guaramacal 

road; ~2,450 m.

Lepanthes wageneri Rchb. f., Bonplandia (Hannover) 3: 70. 

1855; Dunsterville and Garay, Venez. Orchids Ill. 1: 190–

191. 1959; Foldats, Fl. Venez. 15(2): 519–521, fig. 388. 

1970; Romero and Carnevali, Orchids Venez., 2nd ed., 2: 

440. 2000; Morillo, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 1: 207, fig. 70. 2011.

Lepanthes barbae auct., non Schltr.; Foldats, Fl. Venez. 15(2): 463–464, fig. 

465. 1970, pro parte; Ortega et al., BioLlania 5: 47. 1987.

MAP 187. Lepanthes glochidea occurrence in Venezuela. MAP 188. Lepanthes scolex occurrence in Venezuela.
166 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Lepanthes decipiens auct., non Ames & C. Schweinf.; Dunsterville and Garay, Orchids Venez., ed. 1, 2: 387. 1979.

Epiphytic herbs, 2–7.5 cm tall. Leaf blades ovate to broadly 

elliptic or ovate-elliptic, 1.5–2.5 cm × 6–20 mm. Flowers not 

resupinate. Sepals yellow-brown; dorsal sepal 2.3–4 mm long, 

ovate, apices acute. Petals and labellum orange.

Found in South America (Colombia, Venezuela, Ecuador, and Bolivia). In Venezuela, occurs in the Andes (Mérida, 

Táchira, and Trujillo) and in the Cordillera de la Costa (Aragua, 

Distrito Federal, Falcón, Miranda, Sucre, and Yaracuy; Map 

190). Park records are from the north slope of Guaramacal between El Cafenol and Fila de Los Recostaderos and along the 

Boconó–Guaramacal road; 1,800–2,200 m.

The material collected in the Guaramacal area is included 

here in a broadly conceived Lepanthes wageneri (fide Luer, 

1996: 180–181, fig. 260). This concept includes a series of 

populations found in the Venezuelan Andes, the Cordillera de 

la Costa, and possibly Colombia. These populations are characterized by plants of small or intermediate size; dense, abbreviated inflorescences with imbricate, pubescent or hispid floral 

bracts; and elongate floral pedicels. There is, however, some 

diversity in petal and labellum shape. Within the L. wageneri 

complex, the Guaramacal populations are characterized by 

their small vegetative size, thickly succulent leaves, and short 

pedicels. The populations could represent an undescribed species that we hesitate to describe until a revision of the complex 

is completed.

Lepanthes sp. A

Epiphytic herbs, 5–9 cm tall. Leaf blades suborbicular, 

2–2.5 × 2–2.5 mm. Flowers not resupinate, color unknown. Dorsal sepal 3.5–4 mm long, elliptic, apices acuminate.

Evidently endemic to the Andes of Venezuela (Trujillo; Map 

191). Cloud forest along the Boconó–Guaramacal road; 1,800 m.

This appears to be an undescribed species that presently is 

known only from the park.

Lepanthes sp. B

Tiny, epiphytic herbs, 4–6.5 cm tall (including the inflorescences). Leaf blades broadly elliptic or broadly obovate to 

MAP 190. Lepanthes wageneri occurrence in Venezuela.

MAP 189. Lepanthes vareschii occurrence in Venezuela.

MAP 191. Lepanthes sp. A occurrence in Venezuela.
NUMBER 100 • 167

suborbicular, 10–20 mm long. Inflorescences 25–35 mm long. 

Sepals 2.3–2.5 mm long, reddish-purple.

Evidently endemic to the Andes of Venezuela (Portuguesa; 

Map 192). La Divisoria de la Concepción; 1,700 m.

This appears to be an undescribed species that is known 

from a single collection in fruit made in the park.

Lepanthopsis (Cogn.) Ames

Lepanthopsis (Cogn.) Ames, Bot. Mus. Leafl. 1(9): 3. 1933. Pleurothallis

sect. Lepanthopsis Cogn., in Martius, Fl. Bras. 3(4): 591. 1896.

Expedicula Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 103: 308. 2005.

Epiphytic to rarely lithophytic herbs; small to medium in 

size; caespitose. Stems erect, arching or pendulous, thin, terete, 1-leaved apically, enclosed by a series of tubular, ribbed, 

± imbricate sheaths with oblique, dilated ostia or mouths (i.e., 

lepanthiform sheaths), ribs and margins of ostia visibly or 

microscopically ciliate or scabrous; stems often apically proliferous and rooting at the base and apex. Leaves articulate, 

conduplicate; leaf blades elliptic, narrowly elliptic to ovate or 

obovate, bases rounded or attenuate, apices obtuse to acuminate, flat or ± concave, subsessile or short-pseudopetiolate. Inflorescences terminal, 1 to several, originating from near stem 

apex, with an annulus, racemose, 1-flowered (our species) or 

many-flowered, shorter to much longer than subtending leaves; 

peduncles short or elongate; rachises straight or fractiflex. 

Flowers resupinate, minute to small, opening widely, often 

simultaneous or opening in slow succession and then usually 

several to many flowers open simultaneously, bronzy-green, hyaline, yellowish to deep wine-purple. Sepals usually acuminate; 

dorsal sepal usually 3-nerved; lateral sepals free to variously 

connate, similar to the dorsal sepal, often falcate or oblique, 

usually 1-nerved. Petals often shorter and narrower than sepals, 1-nerved, membranous. Labellum usually entire, rarely 

3-lobed, often concave and somewhat concealing the column, 

generally much shorter than sepals; disk callose or ecallose, attached to column base; column short and thick, apically dilated, footless; anther terminal, operculate, incumbent; pollinia 

2, waxy, subpyriform, attached to a small viscidium; rostellum 

transversal; stigma apical, 2-lobed, continuous under rostellar 

flap. Ovary articulate, minutely pedicellate, smooth to costate. 

Capsules ovoid, obovoid or ellipsoid. [Epidendroideae: Epidendreae: Pleurothallidinae.]

A subtropical and tropical American genus of ~50 species 

found in North America (USA), Mexico, Central America, the 

Greater Antilles, and South America (Colombia, Venezuela, 

Guyana, French Guiana, Ecuador, Peru, Bolivia, and Brazil). 

Species diversity is greatest in the Greater Antilles and the Andes. 

Eight species occur in Venezuela.

References. Luer (1986a, 1991); Pridgeon et al. 

(2001b, 2005).

The monophyly of Lepanthopsis has not been established 

yet on a molecular basis, and its relationship to other lepanthiform pleurothallids is not well resolved. Pridgeon et al. (2001b), 

who examined only one species of Lepanthopsis, noted that 

it resolved as part of a larger clade with Frondaria Luer and 

Trichosalpinx.

♦ Lepanthopsis apoda (Garay & Dunst.) Luer, Selbyana 7: 100. 

1982; Romero and Carnevali, Orchids Venez., 2nd ed., 2: 

442. 2000; Fernández, Orquídeas Nat. Táchira 124. 2003; 

Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 1: 209–210, fig. 72, foto 36. 2011. Pleurothallis apoda

MAP 192. Lepanthes sp. B occurrence in Venezuela.

MAP 193. Lepanthopsis apoda occurrence in Venezuela.
168 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Garay & Dunst., in Dunsterville and Garay, Venez. Orchids 

Ill. 3: 246–247. 1965; Foldats, Fl. Venez. 15(2): 203–204, 

fig. 254A. 1970; Ortega et al., BioLlania 5: 49. 1987. Trichosalpinx apoda (Garay & Dunst.) Luer, Phytologia 54: 394. 

1983. Expedicula apoda (Garay & Dunst.) Luer, Monogr. 

Syst. Bot. Missouri Bot. Gard. 103: 308. 2005.

Epiphytic herbs; stems 2–4.5 cm tall. Leaf blades narrowly 

elliptic, 2–2.8 cm × 3.5–5 mm. Sepals yellow to light red-orange; 

dorsal sepal 5–7 mm long. Petals yellow to light red-orange. Labellum orange.

Found in South America (Colombia, Venezuela, Ecuador, 

and Bolivia). In Venezuela, known only from the Andes (Táchira 

and Trujillo; Map 193). Rare or overlooked in dwarf cloud forest below the summit of Guaramacal; 2,700–2,850 m.

In some respects this species appears to be intermediate 

between Lepanthopsis and Trichosalpinx. Its 1-flowered inflorescence, however, is exceptional among the three lepanthiformsheathed genera (Lepanthes, Lepanthopsis, and Trichosalpinx) 

found in Guaramacal. This inflorescence character led Luer 

(2005) to create the genus Expedicula to accommodate our species and a second species, also with a 1-flowered inflorescence, 

from Colombia. Nonetheless, phenetically, our species still seems 

to be better placed in Lepanthopsis.

Lindleyalis Luer

Lindleyalis Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95: 258. 2004.

Epiphytic, rarely lithophytic or terrestrial herbs; erect or patent; relatively robust (in comparison to other Pleurothallidinae). 

Rhizomes short, creeping. Stems terete, erect, straight or somewhat recurved, covered with a remote tubular, appressed central 

sheath and several smaller sheaths at the base; unifoliate apically. 

Leaves articulate; leaf blades elliptic or oblong to ovate-oblong, 

coriaceous. Inflorescences few-flowered racemes or reduced to a 

fascicle of successive flowers, emerging from the apex of the stem 

with an annulus that is very close to the leaf abscission zone; 

racemes usually patent, nutant or pendulous; peduncles with few 

sheaths or remote bracts. Flowers long-pedicellate, resupinate, 

relatively large and showy (in comparison to other Pleurothallidinae), relatively long-lived; perianth segments translucent or 

hyaline, nerves red, purple or dark burgundy. Sepals slightly exserted, ringent, elliptic, narrowly elliptic or lanceolate, acute or 

long-acuminate; lateral sepals fused into a synsepal ± similar to 

dorsal sepal in shape and size, sometimes wider. Petals elliptic 

basally, then prolonged in linear tails 2 or 3× longer than the 

basal portion. Labellum ± adnate to the basal portion of the column or spine or sometimes articulate with it, subentire or with 

basal lobes ± hooked, porrect, central lobe much larger than lateral lobes, elliptic to broadly elliptic, obtuse to acute, with dark 

burgundy veins, glabrous or conspicuously pubescent toward the 

apex; column short, robust, columnar foot pedestal-like; anther 

subventral beneath laminar hood; pollinia 2 on a small hamulus and a tiny, subspherical viscidium; stigma ventral, oblong to 

subquadrate. [Epidendroideae: Epidendreae: Pleurothallidinae.]

A neotropical genus of 8 species found in Central America 

and South America (Colombia, Venezuela, Ecuador, Peru, and 

Bolivia); most diverse in Colombia and Venezuela. Four species 

are found in Venezuela.

The recognition of this genus is based solely on morphology. 

No molecular data have been published that would either support or refute the hypothesis that these species form a distinct, 

monophyletic lineage.

Lindleyalis glossopogon (Rchb. f.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95: 258. 2004. Pleurothallis glossopogon

Rchb. f., Bonplandia (Hannover) 3: 71. 1855; Morillo, in 

Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 250–

251, fig. 95, foto 46. 2011.

Pleurothallis biserrula Rchb. f., Bonplandia (Hannover) 3: 71. 1855; Dunsterville and Garay, Venez. Orchids Ill. 3: 248–249. 1965; Foldats, Fl. 

Venez. 15(2): 213–215, fig. 258. 1970; Ortega et al., BioLlania 5: 49. 

1987; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 757. 2000; 

Dorr et al., Contr. U.S. Natl. Herb. 40: 52. 2000 [2001]; Fernández, 

Orquídeas Nat. Táchira 175. 2003.

Epiphytic herbs, to 45 cm long (usually shorter). Leaf blades 

oblong to ovate-oblong, 7–15 × 1.8–5 cm. Inflorescences arching. 

Flowers large and showy. Sepals translucent in basal 1⁄3 (throughout the basal 1⁄2 in dorsal sepal), otherwise opaque pale burgundy, 

nerves dark burgundy; dorsal sepal 3–6.7 cm long. Petals white; 

labellum opaque burgundy, apex pale greenish-yellow.

Found in South America (Colombia, Venezuela, and Ecuador). In Venezuela, known only from the Andes (Mérida, 

Táchira, and Trujillo; Map 194). Collected along the Boconó–

Guaramacal road and on the trail from Casa Vicuyal to Páramo 

de Vicuyal; (1,850–)2,150–2,800 m.

MAP 194. Lindleyalis glossopogon occurrence in Venezuela.
NUMBER 100 • 169

Lockhartia Hook.

Lockhartia Hook., Bot. Mag. 54: t. 2715. 1827.

Neobennettia Senghas, J. Orchideenfreund 8: 364. 2001.

Epiphytic herbs; small to medium-sized; erect to pendulous; 

caespitose. Stems elongate, subterete to flattened, wholly covered by short, imbricating, laterally compressed sheaths. Leaves 

laterally compressed, narrowly oblong to narrowly or broadly 

triangular, sometimes dorsally keeled, bases imbricate and articulate with their sheaths, apices acute to rounded. Inflorescences 

originating from the leaf axils, usually subapically, frequently 

from the middle or lower internodes of the stem, 1 to many simultaneously, 1- to many-flowered, when many-flowered, short, 

successively to simultaneously few-flowered racemes, rarely 

short, simultaneously flowered lax panicles; bracts of rachis and 

peduncle usually ± foliaceous, flat to concave, relatively conspicuous. Flowers resupinate or not, inconspicuous to relatively 

showy, usually yellow with purple or maroon tinges or spots or 

totally white; floral bracts usually ± foliaceous, flat to concave, 

relatively conspicuous; perianth segments membranous or subfleshy, widely spreading. Petals and sepals similar, sometimes the 

petals somewhat wider, usually elliptic or oblong, rarely ovate 

or obovate, acute, rounded or truncate. Labellum usually flat 

to convex, entire or 3-lobed, when lobed, usually with a pair of 

erect, antrorse to retrorse, short, slender basal lobes, frequently 

± pandurate; apex obtuse to deeply emarginate; disk ecallose 

or variously callose, the upper surface of labellum glabrous to 

variously pubescent; column relatively short, subparallel with 

the labellum forming an obtuse to acute angle with it, laterally 

produced in a pair of well-developed wings; anther apical to ventral, operculate, incumbent; pollinarium consisting of a small 

viscidium, a short or elongate hyaline tegula and 2 obpyriform 

pollinia; rostellum transverse; stigma ventral or subapical. Ovary 

terete. Capsules almost globular. [Epidendroideae: Cymbidieae: 

Oncidiinae.]

A neotropical genus of ~30 species found in Mexico, Central 

America, and South America (Colombia, Venezuela, Trinidad 

and Tobago, the Guianas, Ecuador, Peru, Bolivia, and Brazil). 

Seven species are known from Venezuela.

References. Lambert (1977); Neubig et al. (2012); 

Pridgeon et al. (2009); Senghas (1995b).

Lockhartia chocoensis Kraenzl., in Engler, Pflanzenr. IV (Heft 

83): 19, t. 2, fig. E. 1923; Dunsterville and Garay, Venez. 

Orchids Ill. 6: 220. 1976; Romero and Carnevali, Orchids 

Venez., 2nd ed., 2: 456. 2000 (“chacoënsis”); Fernández, 

Orquídeas Nat. Táchira 125. 2003 (“chacoënsis”).

Epiphytic herbs, erect or ± pendulous, 20–27 cm tall. Leaves 

1.5–2 cm long. Flowers yellow with purple areas. Sepals 5–7 mm 

long.

Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, Ecuador, and Peru). In 

Venezuela, known only from the Andes (Táchira and Trujillo) 

(Map 195). Found once in the karst area above the Río Amarillo 

near the lower limits of the southern boundary of the park; 

~1,200–1,400 m.

Malaxis Sol. ex Sw.

Malaxis Sol. ex Sw., Prodr. 8, 119. 1788.

Tamayorkis Szlach., Fragm. Florist. Geobot., Suppl. 3: 121. 1995.

Terrestrial or rarely epiphytic herbs. Rhizomes usually 

well developed and creeping or very short, prostrate or ascending to erect. Stems usually thickened into a pseudobulb or not; 

pseudobulbs narrowly ovoid, pyriform to subcylindrical, erect, 

covered by sheaths, with 1−3 apical or subapical leaves, rarely 

distichously leaved. Leaves herbaceous or subplicate, not articulate; leaf blades usually diverging and appearing subopposite, 

bases contracted into a long sheath-like petiole, membranous to 

subfleshy; petioles clasping the basal part of the peduncle and 

usually the entire pseudobulb. Inflorescences terminal, racemose 

or (more rarely) spicate, emerging from the pseudostem formed 

by leaf petioles; peduncles terete or ± compressed, usually much 

longer than rachises; rachises often very short and flowers arranged in subumbels; floral bracts inconspicuous. Flowers small 

or minute, usually greenish or yellowish, usually not resupinate 

and presented horizontally; perianth segments usually widely 

spreading, subfleshy. Sepals free or lateral sepals connate below 

the basal 1⁄2, subequal. Petals usually narrower than the sepals, 

often filiform, sometimes twisted. Labellum sessile, not articulate 

and rigidly adnate to the column, fleshier and broader than the 

other perianth segments, entire or 2- or 3-lobed, base concave, 

rounded, cordate, auriculate or sagittate, margins entire or lacerate; column very short, erect, terete; clinandrium conspicuous, 

membranous, oblique; anther terminal, erect, 2-locular; pollinia 

MAP 195. Lockhartia chocoensis occurrence in Venezuela.
170 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

4 in 2 subequal pairs, waxy; rostellum erect; stigma ventral 

to subterminal. Capsules ovoid to ellipsoid. [Epidendroideae: 

Malaxideae.]

Malaxis s.l. is a cosmopolitan genus of 250−300 species, 

with its greatest species diversity in Asia and Oceania and secondary centers of diversity in Mexico and the Andes. In the 

Americas, the genus is found in North America (Canada and 

USA), Mexico, Central America, the West Indies, and South 

America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, 

and Argentina). Approximately 15 species occur in Venezuela.

References. Cameron (2005); Pridgeon et al. (2005).

The species of Malaxis are fundamentally shade loving but 

very variable in terms of their habitat and ecological preferences. 

The two species found in Guaramacal have a pair of subopposite 

leaves and subumbellate inflorescences.

Molecular data indicate that Malaxis s.l. is polyphyletic (Cameron, 2005; Pridgeon et al., 2005), and the genus is likely to be broken up into seven or more genera. It appears likely that our species 

will remain in Malaxis s. str., a large clade of predominantly American taxa. Preliminary data (Pridgeon et al., 2005) also indicate that 

in addition to this large predominantly American clade, there are 

temperate Eurasian–North American, African, and Asian clades.

KEY TO THE SPECIES OF MALAXIS

1a. Labellum apex 3–5-toothed, oblong-obovate in general outline; petals normally straight . . . . . . . . . . . . . . . . M. licatae

1b. Labellum apex rounded or apiculate, ovate-suborbicular in general outline; petals normally spiraled (i.e., twisted like a 

corkscrew) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. nidiae

♦ Malaxis licatae Carnevali & I. Ramírez, Harvard Pap. Bot. 3: 

244, fig. 3. 1998 (“licatiae”).

Malaxis sp. A; Dorr et al., Contr. U.S. Natl. Herb. 40: 51. 2000 [2001].

Terrestrial herbs, 14–25 cm tall. Leaves 2, 4.5–15 × 2.5–7.5 

cm. Flowers green. Dorsal sepal 4–4.5 mm long. Labellum 4–4.5 

mm long.

Endemic to the Cordillera de Mérida in Venezuela (Portuguesa 

and Trujillo; Map 196). Cloud forest near La Divisoria de la Concepción and on the north slope of Guaramacal; 1,700–2,350 m.

Malaxis licatae belongs to the taxonomically difficult complex of species surrounding M. excavata (Lindl.) Kuntze. The latter name has been applied indiscriminately to a large number of 

taxa that have superficially similar labella. This group of species 

is characterized by having apically 3-toothed labella. Malaxis excavata is known with certainty only from Mexico, whereas many 

of the other species in the group are found in southern Central 

America and in the Andes. This species complex merits further 

study.

♦ Malaxis nidiae Carnevali & I. Ramírez, Harvard Pap. Bot. 3: 

246, fig. 4. 1998; Romero and Carnevali, Orchids Venez., 

2nd ed., 2: 482. 2000; Fernández, Orquídeas Nat. Táchira 

128. 2003.

Malaxis parthonii auct., non C. Morren; Foldats, Fl. Venez. 15(1): 439–441, 

fig. 164. 1969, pro parte (i.e., fig. 164, the lower margin depicting the 

labellum in the left of the drawing).

MAP 196. Malaxis licatae occurrence in Venezuela. MAP 197. Malaxis nidiae occurrence in Venezuela.
NUMBER 100 • 171

Terrestrial herbs, 13–24 cm tall. Leaves 2, 4.5–7 × 2.5–4 

cm. Flowers green to brown. Dorsal sepal 3 mm long. Labellum 

3 mm long.

Endemic to the Andes of Venezuela (Táchira and Trujillo; 

Map 197). Cloud forest on the north slope of Guaramacal; 

2,100 m.

Masdevallia Ruiz & Pav.

Masdevallia Ruiz & Pav., Fl. Peruv. Prodr. 122. 1794.

Luerella Braas, Orchidee (Hamburg) 30: 108. 1979.

Rodrigoa Braas, Orchidee (Hamburg) 30: 203. 1979.

Portillia Königer, Arcula 6: 154. 1996.

Masdevallia subgen. Fissia Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 77: 

10. 2000. Fissia (Luer) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 

105: 9. 2006.

Jostia Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 79: 2. 2000.

Actinopetala Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 3. 2006.

Alaticaulia Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 4. 2006.

Buccella Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 7. 2006.

Byrsella Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 7. 2006.

Luzama Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 10. 2006.

Megema Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 11. 2006.

Petalodon Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 11. 2006.

Regalia Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 12. 2006.

Reichantha Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 13. 2006.

Spectaculum Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 14. 2006.

Spilotantha Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 15. 2006.

Sterptoura Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 16. 2006.

Triotosiphon Schltr. ex Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 

16. 2006.

Zahleria Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 17. 2006.

Epiphytic, lithophytic or humicolous herbs; very small 

and inconspicuous to large and showy; erect or patent, rarely 

pendulous; usually caespitose but with some species creeping. 

Stems unifoliate, short to virtually absent compared to the length 

of the leaf, rarely slightly elongate, usually terete, more rarely 

angled, covered with 1 to several scarious sheaths, sometimes 

pustular or dyed black. Leaves with duplicative prefoliation, 

flat to rarely subterete, articulate; leaf blades usually narrowly 

obovate-elliptic but often elliptic to linear-elliptic, sessile to 

long-petiolate, leathery or fleshy, petiolate. Inflorescences emerging from an annulus below the abscission zone (leaf articulation), short- to long-pedunculate, erect, patent to pendulous, 

1- to many-flowered, rarely racemose with several flowers opening simultaneously; peduncles terete or triquetrous, sometimes 

basally triquetrous and apically terete, more rarely somewhat 

laterally compressed, covered with 1–4 sheaths; rachises very 

short to long, usually much shorter than peduncles. Flowers usually resupinate, small or minute to large and showy, short- to 

long-pedicellate. Sepals variously connate, forming a basal cup, 

rarely nearly free, lateral sepals usually forming a conspicuous 

synsepal, apices caudate, color variable but often the inside face 

a different color than the outside one. Petals much smaller than 

sepals and usually enclosed in the sepaline cup, free, flanking the 

column and labellum, usually oblong to oblong-elliptic, apically 

lobed, toothed to rarely entire, with a callus on the margin that 

is usually developed into a ± conspicuous tooth, fleshy. Labellum small, parallel to the column, oblong to elliptic or obovate, 

± fleshy, articulate with a curving extension of the apex of the columnar foot, often contracted near or above the middle and then 

with an apical lobe and basal blade; column terete or semiterete, 

enclosed in the sepaline cup, straight or curved, winged or not; 

anther ventral or subterminal, operculate, incumbent, 1-locular; 

clinandrium entire or variously toothed; pollinia 2, ovoid, waxy, 

attached to a small viscidium; rostellum transverse; stigma 

ventral. Ovary 3-valvate, articulate with pedicel, smooth, verruculose to rugulose. Capsules ellipsoid. [Epidendroideae: Epidendreae: Pleurothallidinae.]

A neotropical genus of ~400 species found in Mexico, Central America, and South America (Colombia, Venezuela, the 

Guianas, Ecuador, Peru, Bolivia, and Brazil). The genus is most 

diverse in the Andes, especially in Colombia, Ecuador, and Peru. 

Approximately 55 species occur in Venezuela.

References. Abele (2007); Luer (1986c, 2000a, 

2000b, 2002a, 2006); Pridgeon and Chase (2001); Pridgeon et 

al. (2001b, 2005).

Masdevallia is a well-supported monophyletic genus most 

closely related to Dracula Luer, Trisetella Luer, and Porroglossum Schltr. (Pridgeon and Chase, 2001; Pridgeon et al., 2001b; 

Abele, 2007). The many segregate genera proposed by Luer 

(2006) scarcely merit mention.

KEY TO THE SPECIES OF MASDEVALLIA

1a. Inflorescences conspicuously longer than the leaves, with several simultaneous flowers; mature leaf blades 15–25 × 3.5–5 

cm; synsepal purplish in the apical 1⁄2, bright yellow in the basal 1⁄2, sepal tails pale purple . . . . . . . . . . . . . . M. schlimii

1b. Inflorescences slightly longer to conspicuously shorter than the leaves when 1-flowered; mature leaf blades to 11 × 1.5 

cm; synsepal the same color as the sepal tails, white, green or yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Stems and petioles ± black; inflorescences erect, longer than the petioles; flowers white with yellow sepal tails; sepal 

tube narrowly cylindrical, longer than wide, sepal tails linear; labellum simple; petals truncate or obtuse, apically lobed

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. tubulosa

2b. Stems and petioles green; inflorescences pendulous, shorter than the petioles; flowers including sepal tails sulfurousyellow with greenish-yellow veins; sepal tube campanulate, as wide as long, sepal tails triangular; labellum with a 

well developed apical lobe; petals acute, simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. gerlachii
172 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

♦ Masdevallia gerlachii Königer, Arcula 2: 38, t. p. 39. 1994.

Masdevallia macroglossa auct., non Rchb. f.; Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 82: 323, figs. 162, 163. 2000.

Masdevallia civilis auct., non Rchb. f. & Warsz.; Foldats, Fl. Venez. 15(3): 

26–28, fig. 393. 1970, pro parte.

Epiphytic herbs, to 15 cm tall. Leaf blades oblong, 7–11 

cm long, obtuse. Inflorescences 1-flowered, pendulous. Flowers 

small, to 16 mm long, 2 cm in diameter.

Endemic to the Cordillera de Mérida in Venezuela (Trujillo; 

Map 198). Known only from the “Páramo de Guaramacal” 

without a precise locality, but certainly from the Trujillo and not 

the Portuguesa portion of the park; 1,800 m.

Masdevallia gerlachii has been confused with M. civilis

Rchb. f. & Warsz., a species evidently endemic to Peru, but the 

inflorescence is pendulous and the labellum is clearly lobed in the 

former as opposed to erect and simple in the latter.

We are not altogether convinced that Masdevallia macroglossa Rchb. f., described from the Cordillera de la Costa, is the 

same as M. gerlachii, described from the Andes, but examination 

of their types could prove us to be wrong.

Masdevallia schlimii Linden ex Lindl., Orchid. Linden. 5. 1846; 

Dunsterville and Garay, Venez. Orchids Ill. 3: 170–171. 

1965; Romero and Carnevali, Orchids Venez., 2nd ed., 

2: 503. 2000; Fernández, Orquídeas Nat. Táchira 136. 

2003; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 216–217, fig. 76, foto 38. 2011. Alaticaulia 

schlimii (Linden ex Lindl.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 6. 2006.

Epiphytic herbs, to 40 cm tall. Leaf blades obovate-elliptic, 

10–21 cm long, obtuse. Inflorescences 4–6-flowered, erect. Flowers showy, to 2 cm long, 10 cm in diameter.

Found in South America (Colombia and Venezuela). In Venezuela, known only from the Andes (Mérida, Táchira, and Trujillo; Map 199). Without precise locality; 2,500 m.

There is some doubt as to whether or not this species has 

been collected within the boundaries of Guaramacal National 

Park. The locality of our voucher (Renz 4651, RENZ) is vague, 

and the specimen may have been collected to the northwest of 

the park along the Boconó–Trujillo road.

Masdevallia tubulosa Lindl., Orchid. Linden. 4. 1846; Foldats, Fl. 

Venez. 15(3): 59–61, fig. 406. 1970; Dunsterville and Garay, 

Venez. Orchids Ill. 5: 178–179. 1972; Romero and Carnevali, 

Orchids Venez., 2nd ed., 2: 511. 2000; Fernández, Orquídeas 

Nat. Táchira 137. 2003; Morillo, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 1: 219–220, fig. 792. 2011.

Epiphytic herbs, 6.5–11 cm tall. Leaf blades oblongoblanceolate, 4–12 cm long, acute or subacute. Inflorescences 

1-flowered, erect. Flowers 3–7 cm long, to 10 cm in diameter, 

tubular, white.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, reported from the Andes (Mérida, 

Táchira, and Trujillo) and the Cordillera de la Costa (Aragua 

and Distrito Federal; Map 200). Collected on the north slope of 

Guaramacal; 1,850–2,200 m.

Maxillaria Ruiz & Pav.

Maxillaria Ruiz & Pav., Fl. Peruv. Prodr. 116. 1794.

Calawaya Szlach. & Sitko, in Szlachetko et al., Biodivers. Res. Conservation 

25: 23. 2012.

Pseudocymbidium Szlach. & Sitko, in Szlachetko et al., Biodivers. Res. Conservation 25: 33. 2012.

MAP 198. Masdevallia gerlachii occurrence in Venezuela. MAP 199. Masdevallia schlimii occurrence in Venezuela.
NUMBER 100 • 173

Epiphytic, lithophytic or rarely terrestrial herbs; caespitose to climbing or creeping. Rhizomes very short to long and 

creeping or climbing; pseudobulbs present (our species) or absent, spherical, ovoid or oblongoid, with or without distichous 

and ± imbricate sheaths, innermost sheaths sometimes with foliar blades. Leaves conduplicate to semiterete, articulate with 

their sheaths or with the apex of the pseudobulb; pseudobulbs 

1-foliate (our species) or leaves distributed distichously along the 

stems; leaf blades oblong, oblong-elliptic, linear or subterete to 

broadly elliptic to suborbicular, apices irregularly oblique, bases 

attenuate, submembranous, leathery to succulent, sessile or petiolate. Inflorescences 1-flowered, 1 or several flowers produced 

simultaneously, originating from the bases of the pseudobulbs or 

from the axils of the leaf sheaths, covered with scarious sheaths. 

Flowers inconspicuous to large and showy, usually resupinate, 

variously colored; floral bracts usually inconspicuous, shorter 

than the ovary; perianth free, parallel to the column to widely 

expanded, submembranous or fleshy. Sepals usually longer and 

wider than petals; lateral sepals oblique. Petals subparallel to the 

column. Labellum fleshy, articulate or not to the column foot, 

usually 3-lobed, central lobe usually larger than the lateral lobes; 

labellar disk usually equipped with several calli or longitudinal 

ridges that are usually made of wax; other calli are located on 

the central lobe; column usually long, cylindrical or semicylindrical, straight or recurved; anther apical, operculate, incumbent; 

clinandrium often provided with teeth or marginal cilia; pollinia 

4 in 2 unequal pairs, superimposed dorsoventrally; viscidium 

well developed, hippocrepiform, stipes lunate or absent sometimes elongate; rostellum transverse; stigma solitary, ventral, 

transverse to ovoid. Ovary cylindrical to 3-angled, pedicellate. 

Capsules ellipsoid or subspherical, sometimes 3-angled, surface 

usually smooth, laterally dehiscent, marcescent perianth often at 

apex. [Epidendroideae: Cymbidieae: Maxillariinae.]

A neotropical genus of 200–250 species found in Mexico, 

Central America, and South America (Colombia, Venezuela, the 

Guianas, Ecuador, Peru, Bolivia, Brazil, and Argentina); most 

diverse in the Andes and southern Central America. Approximately 100(or fewer) species are found in Venezuela.

References. Blanco et al. (2007); Christenson 

(2002a, 2002b); Pridgeon et al. (2009); Senghas (2002); Whitten 

et al. (2007).

Maxillaria s.l. is grossly polyphyletic (Blanco et al., 2007; Whitten et al., 2007), which presents a number of taxonomic challenges. 

Most fundamentally, either one recognizes an extremely large and 

variable genus within the “core Maxillariinae,” or one divides Maxillaria s.l. into many smaller, monophyletic clades. We have attempted 

to follow the latter course by recognizing not only Maxillaria s. str. 

but also Camaridium and Ornithidium R. Br. None of the other segregate genera of a much-reduced Maxillaria are known to occur in 

Guaramacal. The many generic segregates proposed by Szlachetko 

et al. (2012) seem to be based on a few homoplasious morphological characters and/or incomplete molecular sampling, and we see no 

value in adopting their concepts that diverge from those based on the 

published molecular phylogeny of Whitten et al. (2007).

KEY TO THE SPECIES OF MAXILLARIA

1a. Pseudobulbs widely separated, conical-fusiform, completely covered when young by several imbricate sheaths of which 

the innermost 1 or 2 have foliar blades; leaf blades ≤15 cm long, 15–20× longer than wide; sepals ≤2.5 cm long . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. meridensis

1b. Pseudobulbs clustered on short rhizomes, not fully covered by their sheaths, which lack foliar blades; leaf blades usually 

≥15 cm long, 8–15× longer than wide; sepals ≥2.5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Sepals 9–10 cm long, linear-elliptic; dorsal sepal long and narrowing apically; flowers pale purple or pink with dark 

purple longitudinal striations; callus small, restricted to the basal 1⁄4 of the labellum; pollinia suborbicular . . . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. macrura

2b. Sepals 5–7 cm long, oblong or elliptic; dorsal sepal not narrowing apically or only abruptly in the lower 1

⁄6; flowers yellow, orange or dark purple; callus large, occupying the entire basal 1⁄2 of the labellum; pollinia narrowly elliptic . . . 3

 3a. Flowers deep wine-purple; labellum simple, subquadrate or oblong-subquadrate in general outline; dorsal sepal 

oblong-elliptic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. nigrescens

 3b. Flowers orange-yellow, with white areas; labellum conspicuously 3-lobed, ovate-elliptic in general outline; dorsal 

sepal spathulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. triloris

MAP 200. Masdevallia tubulosa occurrence in Venezuela.
174 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Maxillaria macrura Rchb. f., Linnaea 41: 28. 1877 [1876]; Dunsterville and Garay, Venez. Orchids Ill. 3: 186–187. 1965; 

Foldats, Fl. Venez. 15(4): 476–478, fig. 737. 1970; Romero 

and Carnevali, Orchids Venez., 2nd ed., 2: 556. 2000.

Epiphytic herbs. Leaf blades 15–35 cm long. Inflorescences 

much longer than the pseudopetioles of the leaves. Perianth segments widely spreading, bases pale flesh color suffused with a 

maroon-tan. Sepals 9–10 cm long.

Endemic to the Venezuelan Andes (Mérida, Táchira, and 

Trujillo; Map 201). In cloud forest in Qda. Segovia along the 

Boconó–Guaramacal road; ~2,600 m.

This species is extremely showy. Sterile material can be confused in our area with Maxillaria nigrescens Lindl., but the old 

peduncles of the inflorescences of M. macrura are much longer 

than the pseudopetioles of the leaves.

A yellow-flowered form of this Venezuelan endemic, Maxillaria macrura f. aurea Christenson, was described from material 

flowering in cultivation in England. The exact provenance of this 

form either is unknown or was suppressed by the author.

Maxillaria meridensis Lindl., Orchid. Linden. 19. 1846; Dunsterville and Garay, Venez. Orchids Ill. 2: 206–207. 1961; Foldats, Fl. Venez. 15(4): 483–486, fig. 740. 1970; Romero and 

Carnevali, Orchids Venez., 2nd ed., 2: 561. 2000; Carnevali 

and Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 7: 

440. 2003; Fernández, Orquídeas Nat. Táchira 146. 2003. 

Calawaya meridensis (Lindl.) Szlach. & Sitko, in Szlachetko 

et al., Biodivers. Res. Conservation 25: 24. 2012.

Subterrestrial, lithophytic or epiphytic herbs, erect or creeping, 25–100 cm long. Leaf blades 10–15 cm long. Inflorescences 

solitary in the sheath axils; peduncles 1.5–3 cm long. Perianth 

segments not widely spreading, greenish, yellowish-brown to 

pink tinged or orange-pink. Dorsal sepal 1.8–2.5 cm long.

Found in Central America (Costa Rica and Panama), the 

Lesser Antilles (Guadeloupe), and South America (Colombia, 

Venezuela, Guyana, Ecuador, Peru, Bolivia, and Brazil). In Venezuela, this species is reported from Amazonas, Aragua, Bolívar, 

Delta Amacuro, Mérida, Miranda, Táchira, Trujillo, and Yaracuy (Map 202). The sole collection made in the park was from 

a disturbed area below the Páramo del Pumar; ~2,600 m.

Maxillaria nigrescens Lindl., Orchid. Linden. 20. 1846; Dunsterville and Garay, Venez. Orchids Ill. 3: 188–189. 1965; 

Foldats, Fl. Venez. 15(4): 493–494, fig. 744. 1970; Romero 

and Carnevali, Orchids Venez., 2nd ed., 2: 565. 2000; 

Fernández, Orquídeas Nat. Táchira 147. 2003.

Epiphytic herbs. Leaf blades 25–52 cm long. Inflorescences 

± as long as the pseudopetioles of the leave. Perianth segments 

widely spreading; reddish-brown, maroon or yellow with a dark 

maroon throat. Sepals 2.5–6.7 cm long, widely spreading.

Found in South America (Colombia and Venezuela). In Venezuela, known from the Andes (Mérida, Táchira, and Trujillo) 

and the Cordillera de la Costa (Miranda; Map 203). In the park, 

found in cloud forest in Qda. Segovia on the north slope and 

along the trail between El Campamento (below Cerro El Diablo) 

and El Maciegal on the south slope; 1,800–1,900 m.

In this species, the petals arch downward.

Material from Ecuador and Peru identified as Maxillaria 

nigrescens can be referred to M. calantha Schltr. and is readily 

distinguished by its longer peduncles.

MAP 201. Maxillaria macrura occurrence in Venezuela. MAP 202. Maxillaria meridensis occurrence in Venezuela.
NUMBER 100 • 175

Maxillaria triloris E. Morren, Belgique Hort. 20: 240, t. 13. 

1870; Dunsterville and Garay, Venez. Orchids Ill. 1: 244. 

1959; Foldats, Fl. Venez. 15(4): 538–539, fig. 762. 1970; 

Fernández, Orquídeas Nat. Táchira 153. 2003.

FIGURE 25K

Maxillaria luteograndiflora Hort. ex Dombrain, Fl. Mag. (London) 10: t. 

559. 1871, nom. nud.

Epiphytic herbs, to 70 cm tall. Leaf blades 45–55 cm long. 

Inflorescences shorter than petioles. Perianth segments widely 

spreading. Sepals white basally, yellow in the upper 1⁄2 with some 

purplish-brown in the middle; dorsal sepal 7–7.5 cm long. Petals 

white. Labellum white with yellow in the central lobe and purple 

veins on the labellar disk.

Found in South America (Colombia and Venezuela). In Venezuela, restricted to the Andes (Mérida, Táchira, and Trujillo; 

Map 204). Collected in forest near the lower limits of the southern boundary of the park in the karst area above the Río Amarillo; ~1,200–1,400 m.

Although Maxillaria triloris is very showy, it can be confused with several other species found in Venezuela, including 

M. setigera Lindl. and M. luteoalba Lindl. Unlike our species, 

M. setigera, a species of lower elevations, is smaller vegetatively 

and has strongly compressed and disciform pseudobulbs that are 

proportionally wider and flowers that are smaller with narrower 

sepals to 4.5 cm long. Similarly, M. luteoalba, unlike M. triloris,

has sharply defined petioles and flowers that lack any suggestion 

of brown pigmentation.

Christenson (2004) argued that Maxillaria luteograndiflora

Hort. ex Dombrain is the correct name for M. triloris, but we 

are not convinced by his arguments, and in any case the name 

he adopted is a nomen nudum as it lacks a description or diagnosis. Maxillaria triloris was described from material cultivated 

in Belgium, and although the exact provenance of this material 

was unknown, there was speculation in the protologue that the 

plants originated in Caracas. Christenson (2004) ignored this 

suggestion and argued that material matching what he called 

the “type” (presumably the plate) only occurs in Ecuador and 

Peru, but the distinguishing petal characters that he cited (petals initially straight and parallel to the column and then curving 

outward) cannot be interpreted from the protologue or accompanying plate.

Microchilus C. Presl

Microchilus C. Presl, Rel. Haenk. 1: 94. 1827.

Primarily terrestrial and secondarily subepiphytic herbs; 

sympodial; sympodia consisting of stems that are initially erect 

and then become decumbent and prostrate, basally trailing; 

stems terete, rooting at the internodes, leafy in the middle or 

upper third, sheaths remote. Leaves convolute, spirally arranged 

on the stem, not articulate, blades usually elliptic or lanceolate, obtuse, basally attenuate into a channeled pseudopetiole 

extending to a funnel-shaped sheath that embraces the stem, 

apically acute or acuminate, usually green but often spotted or 

variegated with silvery streaks or purple tints, herbaceous to 

herbaceous-subfleshy. Inflorescences terminal, usually manyflowered spikes; peduncles and rachises glandular-pubescent; 

peduncles usually well developed, covered by 2 or 3 tubular, 

appressed sheaths; floral bracts ± conspicuous, usually elliptic 

to ovate-elliptic, glabrous or pubescent, subequal to ovary in 

MAP 203. Maxillaria nigrescens occurrence in Venezuela. MAP 204. Maxillaria triloris occurrence in Venezuela.
176 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

length. Flowers resupinate, small, usually rather thin-textured; 

sepals free, subequal; dorsal sepal concave to cymbiform, connivent with the petals; lateral sepals connivent to spreading. 

Petals connivent with the dorsal sepal to form a galea or ± 3-

lobed hood, linear to oblong-lanceolate. Labellum attached by 

its margins to the basal portion (1⁄3–1⁄2) of the column, divided 

into hypochile (i.e., basal part of a complex labellum, which 

is divided into 2 or 3 distinct parts) and epichile (i.e., terminal 

part of a complex labellum, which is divided into 2 or 3 distinct 

parts), extended basally into a cylindrical or ellipsoidal spur, 

sometimes conically bilobed, obtuse or subacute, usually with 

low thickenings that extend in veins from the base to the apex of 

the spur; hypochile semitubular, flattened rectangular to broadly 

elliptic, rarely keeled internally, often constricted apically; epichile sessile or with a short claw, often with a keel or papillose 

thickening extending to the opening of the hypochile, entire or 

transversely bilobed, apical lobes perpendicular to the labellar axis or retrorse. Column relatively short, fusiform, basally 

semiterete, sessile or subsessile, somewhat dilated around the 

stigmatic lobes; anthers erect, dorsal, ovoid to lanciform, bilocular; pollinia 4, sectile, clavate-obovoid, attached the length of 

the rostellum that is narrowly clavate-linear, caudicles attached 

to a well-defined viscidium; rostellum elongate, broadly triangular, the remains (after removal of the pollinia) deeply bifid; stigmatic lobes 2, ventral, confluent to connate, transversely elliptic 

or V-shaped. Ovary cylindric, glabrous to (more commonly) pubescent, spirally twisted. Capsules fusiform to obovoid; pedicels 

not elongating. [Orchidoideae: Cranichideae: Goodyerinae.]

A neotropical genus of 45–50 species found in Mexico, Central America, the West Indies, and South America (Colombia, 

Venezuela, Guyana, Suriname, Ecuador, including the Galapagos Islands, Peru, Bolivia, Brazil, Paraguay, Argentina, and Uruguay). Sixteen species occur in Venezuela.

References. Ormerod (2002); Pridgeon et al. (2003).

Species of Microchilus are usually shade loving and found 

in leaf duff.

Microchilus, which often is included in a broadly circumscribed Erythrodes Blume, can be distinguished by its deeply 

bifid rostellum (following removal of the pollinia) and by its 

neotropical distribution.

Microchilus paleaceus (Schltr.) Ormerod, Lindleyana 17: 219. 

2002. Physurus paleaceus Schltr., Repert. Spec. Nov. Regni. 

Veg. Beih. 7: 72. 1920. Erythrodes paleacea (Schltr.) Ames, 

Orchidaceae 7: 75. 1922; Romero and Carnevali, Orchids 

Venez., 2nd ed., 1: 327. 2000; Dorr et al., Contr. U.S. Natl. 

Herb. 40: 50. 2000 [2001]; Carnevali and Ramírez-Morillo, 

in Berry et al., Fl. Venez. Guayana 7: 361. 2003.

Terrestrial humicolous, umbraticolous herbs, 50–100 cm 

tall. Flowers pubescent externally; perianth segments parallel to 

the column. Sepals green or yellow; dorsal sepal 6–8 mm long. 

Petals and labellum white or whitish; spur greenish.

Found in South America (Colombia, Venezuela, Trinidad 

and Tobago, Guyana, Suriname, Ecuador, Peru, and Brazil). 

Widespread in Venezuela (Amazonas, Apure, Aragua, Bolívar, 

Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, 

Monagas, Portuguesa, Sucre, Yaracuy, and Zulia; Map 205). 

In the park, found near La Divisoria de la Concepción; 1,500–

1,800 m.

Myoxanthus Poepp. & Endl.

Myoxanthus Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 50. 1835 [1836].

Epiphytic but sometimes lithophytic or terrestrial herbs or 

suffruticose; inconspicuous to rather large in size; caespitose. 

Rhizomes very short to long and creeping, appressed to substrate or rarely ascending, simple or branched, mostly covered 

with hispidulous sheaths. Stems erect to ascending, 3–50 cm 

tall, subterete, apically 1-foliate, wiry to stout, sometimes distinctly swollen at the base, often apically proliferous, clothed by 

large, papery, usually hispidulous sheaths that often are easily 

shed. Leaves conduplicate, linear to broadly elliptic or oblong, 

thin to rigidly coriaceous. Inflorescences originating without an 

annulus from or near the apex of stem and then 1-flowered, or 

a fascicle of single flowers produced successively or simultaneously. Flowers resupinate, inconspicuous to relatively showy, 

campanulate to widely spreading; perianth segments often 

fleshy, variously pubescent; floral bracts inconspicuous, often 

hispidulous. Sepals free or the laterals variously connate into 

a synsepal, often fleshier and broader than the petals. Petals 

similar to sepals or longer and thinner or much shorter, often 

variously thickened, the apices sometimes developed into osmophores. Labellum articulate to column foot, simple to 3- or 

5-lobed, usually much smaller than the other perianth segments, often convex and recurved, variously callose or ecallose; 

MAP 205. Microchilus paleaceus occurrence in Venezuela.
NUMBER 100 • 177

column well developed, basally produced into a well-developed 

column foot, apically variously toothed or winged; anther ventral to subterminal, incumbent, operculate, sometimes spiculate; clinandrium entire to variously dentate or lacerate; pollinia 

2, attached to a small viscidium; rostellum transverse; stigmatic 

surface ventral. Ovary smooth or hispidulous to echinate. Capsules ovoid, mostly hispidulous to echinate. [Epidendroideae: 

Epidendreae: Pleurothallidinae.]

A neotropical genus of ~40 species found in Mexico, Central America, and South America (Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Bolivia, and Brazil). The greatest 

species diversity is in the Andes; ~12 species occur in Venezuela.

References. Luer (1982, 1992); Pridgeon and 

Chase (2001); Pridgeon et al. (2001b, 2005).

There is strong molecular support for the monophyly of 

Myoxanthus (Pridgeon and Chase, 2001; Pridgeon et al., 2001b). 

Morphologically, most species of Myoxanthus are distinguished 

easily from other pleurothallid orchids by the pubescent to hispidulous sheaths on their stems. This character, however, is not 

exclusive to Myoxanthus.

Myoxanthus reymondii (H. Karst.) Luer, Selbyana 7: 49. 1982; 

Romero and Carnevali, Orchids Venez., 2nd ed., 2: 612. 

2000 (“raymondii”); Carnevali and Ramírez-Morillo, in 

Berry et al., Fl. Venez. Guayana 7: 460. 2003; Fernández, 

Orquídeas Nat. Táchira 157. 2003 (“raymondii”). Duboisia reymondii H. Karst., Allg. Gartenzeitung 15: 394. 1847 

(“Reymondi”). Pleurothallis reymondii (H. Karst.) Rchb. 

f., Ann. Bot. Syst. 3(3): 520. 1852 (“reymondi”); Dunsterville and Garay, Venez. Orchids Ill. 1: 342–343. 1959 

(“raymondii”); Foldats, Fl. Venez. 15(2): 381–383, fig. 329. 

1970 (“reymondi”); Ortega et al., BioLlania 5: 49. 1987 

(“raymondii”).

Epiphytic or rarely lithophytic herbs. Leaf blades 8–20 cm 

long. Sepals spreading, externally purple-brown, internally pale 

or dark honey-brown; dorsal sepal 5–8 mm long. Petals parallel to column, basally pale yellow-brown, apically with dark 

purple-brown osmophores. Labellum yellow with dark brown 

patches.

Found in South America (Colombia, Venezuela, Ecuador, and Peru). Venezuelan records are from the Andes (Lara, 

Mérida, Portuguesa, Táchira, and Trujillo), the Cordillera de la 

Costa (Aragua, Distrito Federal, Falcón, Miranda, and Yaracuy), 

and the Venezuelan Guayana (Bolívar; Map 206). Rare in cloud 

forest along the Boconó–Guaramacal road; 2,100–2,300 m.

Odontoglossum Kunth

Odontoglossum Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 281. 1815 [1816]; 

ibid. [qu.] 1: 350. 1815 [1816].

Symphyglossum Schltr., Orchis 13: 8. 1919, nom. cons.

Epiphytic, lithophytic or subterrestrial herbs; inconspicuous to large and showy; usually erect but sometimes pendulous; 

caespitose to creeping or scandent. Rhizomes abbreviated to 

elongate, prostrate to ascendent, naked or covered with sheaths 

bearing pseudobulbs at variable distances; pseudobulbs heteroblastic, usually ovoid to subspheroidal, sometimes oblong, apically 1–3(–4)-foliate, usually clothed by 1 to several imbricate 

sheaths, the innermost 1–3 of which may have foliar blades. 

Leaves conduplicate, articulate, erect or spreading, coriaceous to 

fleshy, sessile to shortly pseudopetiolate. Inflorescences originating from the base of pseudobulbs or from axils of pseudobulb 

sheaths, usually racemose or variously branched panicles, rarely 

few-flowered, erect, arching or pendulous, sometimes voluble; 

peduncles, rachises, and bracts various. Flowers resupinate, 

medium-sized to large and showy, patent to nodding, usually 

long-lasting; perianth segments spreading to campanulate, flat 

or undulate, mostly white, yellowish or pink, usually spotted, 

striped or blotched with red, purple, maroon, pink or brown, 

sometimes the entire flower purple or maroon. Sepals usually subequal, free or rarely lateral sepals connate in the basal 1⁄3, usually 

ovate-elliptic or elliptic, acute to acuminate, sometimes basally 

clawed. Petals usually similar to the sepals, rarely different. Labellum basally ± parallel to column, then usually deflexed and making an angle of ± 90° to the column, margin entire or denticulate 

to fimbriate, simple or 3-lobed; lateral lobes erect or spreading; 

disk variously callose, the callus denticulate or cristate or composed of several longitudinal ridges or plates, very rarely ecallose; 

column usually elongate and straight, footless, usually with a pair 

of entire or denticulate to lacerate or setiform subapical wings; 

anther terminal, operculate, incumbent, 1-locular or imperfectly 

2-locular; pollinia 2, waxy, mostly ovoid, stipe narrowly oblong, 

viscidium small; clinandrium usually entire, shallow; rostellum 

transverse; stigma ventral. Ovary mostly terete, pedicellate. Capsules various. [Epidendroideae: Cymbidieae: Oncidiinae.]

MAP 206. Myoxanthus reymondii occurrence in Venezuela.
178 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

A South American genus of ~100 species found almost exclusively in the Andes of Colombia, Venezuela, Ecuador, Peru, 

and Bolivia, but with 1 species also in the Guayana Highlands 

(Venezuela and Guyana). Fifteen species occur in Venezuela.

References. Bockemühl (1989); Chase (1997, 

2002); Garay (1970); Neubig et al. (2012); Senghas (1997c).

A revision of Odontoglossum by Bockemühl (1989) settled 

briefly the long-standing debate regarding the distinctiveness of 

Odontoglossum and Oncidium, but with the introduction of molecular characters the debate has been renewed. Oncidium has been 

a catchall genus, and work to delimit a monophyletic Oncidium is 

incomplete; a number of segregate genera have been described or 

resurrected, including Cyrtochilum, Otoglossum (Schltr.) Garay 

& Dunst., and Trichocentrum Poepp. & Endl., which are recognized in this flora. There is a core group of Odontoglossum species 

that are monophyletic, but they are deeply embedded in a broadly 

circumscribed Oncidium (Chase et al., 2008; Neubig et al., 2012; 

Pridgeon et al., 2009). Inasmuch as these species of Odontoglossum also share a similar morphology, geography, and ecology, we 

have chosen to recognize them at the generic level. We appreciate 

that strict adherence to our circumscription implies that Oncidium

will be further subdivided into smaller genera.

KEY TO THE SPECIES OF ODONTOGLOSSUM

1a. Petals and sepals 25–35 × 3–5 mm, narrowly lanceolate, acuminate, 4–5× longer than wide; pseudobulbs 2- or 3-foliate 

with conspicuously thickened internode of the second leaf . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. dormanianum

1b. Petals and sepals 20–26 × 8–11 mm, elliptic or lanceolate, acute, ~2× as long as wide; pseudobulbs 1- or 2-foliate without 

thickened internodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. schillerianum

Odontoglossum dormanianum Rchb. f., Gard. Chron., n.s., 

21: 11. 1884 (“Dormanianum”); Dunsterville and Garay, 

Venez. Orchids Ill. 4: 172–173. 1966 (“dormannianum”); 

Foldats, Fl. Venez. 15(5): 213–214, fig. 840. 1970 (“dormannianum”). Odontoglossum crocidipterum subsp. dormanianum (Rchb. f.) Bockemühl, Odontoglossum, Monogr. 

162. 1989; Dorr et al., Contr. U.S. Natl. Herb. 40: 51–52. 

2000 [2001] (“dormannianum”). Odontoglossum crocidipterum f. dormanianum (Rchb. f.) O. Gruss & M. Wolff, 

Orchid Atlas 244. 2007. Oncidium crocidipterum subsp. 

dormanianum (Rchb. f.) M. W. Chase & N. H. Williams, in 

Chase et al., Lindleyana 21: 24. 2008.

Odontoglossum crocidipterum auct., non Rchb. f.; Romero and Carnevali, 

Orchids Venez., 2nd ed., 2: 655. 2000; Fernández, Orquídeas Nat. 

Táchira 161. 2003.

Odontoglossum sanderianum auct., non Rchb. f.; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 52. 2000 [2001].

Epiphytic herbs; pseudobulbs ~6 × 3 cm. Leaf blades ~25 

× 4 cm. Inflorescences suberect. Sepals and petals white or clear 

pale yellow to brownish-yellow with maroon spots. Labellum 

the same color as perianth but lighter.

Found in South America (Colombia and Venezuela). In Venezuela, found in the Andes (Lara, Mérida, Táchira, and Trujillo) 

and in the Cordillera de la Costa (Sucre; Map 207). In cloud 

forest along the Boconó–Guaramacal road and in sector El Santuario on the south slope of Guaramacal; 1,950–2,700 m.

This species belongs in the taxonomically difficult Odontoglossum constrictum Lindl. complex, which is most diverse in 

Colombia.

Odontoglossum schillerianum Rchb. f., Bonplandia (Hannover) 

2: 12. 1854; Dunsterville and Garay, Venez. Orchids Ill. 3: 

214–215. 1965; Foldats, Fl. Venez. 15(5): 235–237, fig. 

850. 1970; Romero and Carnevali, Orchids Venez., 2nd 

ed., 2: 667. 2000; Fernández, Orquídeas Nat. Táchira 163. 

2003; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 230–231, fig. 83, foto 42. 2011.

Odontoglossum cinnamomeum R. Warner & B. S. Williams, Orchid Album 

3: t. 143. 1884. Oncidium cinnamomeum (R. Warner & B. S. Williams) M. W. Chase & N. H. Williams, in Chase et al., Lindleyana 21: 

22. 2008.

Epiphytic herbs; pseudobulbs 4–7.5 × 2.5–3.5 cm. Leaf 

blades 9–30 × 1.8–3 cm. Inflorescences erect. Sepals yellow with 

brown or purple spots. Labellum white basally.

Endemic to Venezuela, where it is found only in the Andes 

(Lara, Mérida, Táchira, and Trujillo; Map 208). Collected in 

cloud forest along the Boconó–Guaramacal road; 2,450 m.

MAP 207. Odontoglossum dormanianum occurrence in Venezuela.
NUMBER 100 • 179

This species is a member of the Odontoglossum constrictum

complex. In Guaramacal National Park, this species can only 

be confused with the more common O. dormanianum. Odontoglossum schillerianum, however, is easy to recognize by its 

smaller flowers with broader perianth segments.

If one considers Odontoglossum to be a synonym of Oncidium, then the correct name for this species is Oncidium 

cinnamomeum (R. Warner & B. S. Williams) M. W. Chase & 

N. H. Williams because a new combination in Odontoglossum is 

blocked by Odontoglossum schillerianum Rchb. f. (Bonplandia 

(Hannover) 2: 91. 1854).

Oncidium Sw.

Oncidium Sw., Kongl. Vetensk. Acad. Nya Handl. 21: 239. 1800, nom. cons.

Heteranthocidium Szlach. et al., Polish Bot. J. 51: 54. 2006.

Epiphytic or terrestrial, occasionally lithophytic, herbs; 

caespitose to prostrate. Roots thin to thick, white. Rhizomes 

short or long and creeping to scandent. Stems thickened into 

well-developed, clustered, laterally compressed, apically 2- or 

3-foliate pseudobulbs; pseudobulbs partially covered by foliose 

sheaths or not, triangular when sheathless. Leaves conduplicate, 

articulate; leaf blades at the apex of pseudobulbs, flat, flexible 

or rigid, leathery, usually keeled on the back along the midvein. 

Inflorescences lateral, usually originating from base of mature 

pseudobulbs in axils of leaf sheaths, paniculate, occasionally 

racemose, erect or arched; floral bracts triangular. Flowers resupinate, inconspicuous to large and showy, usually yellow with 

brown spots, but in several species entirely white or pink with 

or without brown or purple spots, or entirely coffee-brown, 

membranous to fleshy. Sepals subequal; lateral sepals oblique, 

spreading to reflexed, free or basally connate. Petals mostly 

subequal to sepals. Labellum at the base of the column, entire, 

panduriform or 3-lobed, usually middle lobe larger than lateral 

ones; callus at the base of the labellum, glabrous or papillose, 

rarely pubescent, formed by keels or tubercles, usually a different color than the labellum; column usually thick and short, 

sometimes thin and elongate, without a foot, usually with a pair 

of petaloid processes (wings or projections) on the sides of the 

stigma, usually with a thickening at the base; anther terminal, 

operculate, incumbent, concave, 1- or 2-locular; pollinarium 

consisting of 2 obovoid or oblong-obovoid pollinia, usually yellow or orange, sulcate, with white hyaline laminar or tubular 

stipe, and viscidium usually ovoid, brown; rostellum usually 

short, sometimes elongate, generally triangular; stigmatic cavity 

on the ventral side of the column, entire, ovate to suborbicular 

or triangular, concave, shiny. Ovary terete, sulcate, pedicellate. 

Capsules generally ellipsoid to obovoid. [Epidendroideae: Cymbidieae: Oncidiinae.]

A neotropical genus of ~450 species found in Mexico, Central America, the West Indies, and South America (Colombia, 

Venezuela, the Guianas, Ecuador, Peru, Bolivia, and Brazil); 

most diverse in the Andes. Approximately 38–40 species occur 

in Venezuela.

References. Chase (1986, 1997, 2002); Chase and 

Palmer (1992); Chase et al. (2008, 2009); Garay (1970); Garay 

and Stacy (1974); Neubig et al. (2012); Pridgeon et al. (2009); 

Senghas (1997a); Williams et al. (2001).

Circumscription of Oncidium has been contentious, and 

our treatment of the genus in a more or less traditional sense 

may not prove to be defensible as molecular studies (Chase et 

al., 2008; Pridgeon et al., 2009; Neubig et al., 2012) indicate 

that Odontoglossum and other morphologically distinguishable genera such as Sigmatostalix Rchb. f. are deeply embedded 

within a larger Oncidium clade. Although these segregate genera 

are monophyletic, their position within the phylogenies in the 

molecular studies cited above suggests their recognition would 

require the creation of many new genera. Whether or not that 

is useful taxonomically is beyond the scope of this flora. We do 

accept the exclusion of almost all of the Brazilian species of Oncidium, which are now treated as Gomesa R. Br. (Pridgeon et al., 

2000; Chase et al., 2009).

Many species and hybrids of Oncidium s.l. are grown as 

ornamentals.

Oncidium abortivum Rchb. f., Linnaea 22: 847. 1849; Dunsterville and Garay, Venez. Orchids Ill. 1: 262–263. 1959; Foldats, Fl. Venez. 15(5): 271–274, fig. 858. 1970; Romero and 

Carnevali, Orchids Venez., 2nd ed., 2: 670. 2000. Heteranthocidium abortivum (Rchb. f.) Szlach. et al., Polish Bot. J. 

51: 54. 2006.

Small epiphytic herbs, often growing on twigs, erect; pseudobulbs 2–5 cm in diameter, suborbicular, strongly laterally 

compressed, 1-foliate laterally. Leaf blades oblong to oblonglanceolate, 5–15 cm long. Inflorescences with fertile and abortive 

MAP 208. Odontoglossum schillerianum occurrence in Venezuela.
180 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

flowers. Fertile flowers yellow with brownish-red spots and lines. 

Labellum proportionally large, 1.5–2 cm wide.

Found in Central America (Panama) and South America 

(Colombia, Venezuela, Ecuador, and Peru). In Venezuela, this 

species is known from the Andes (Portuguesa and Táchira) and 

the Cordillera de la Costa (Aragua, Distrito Federal, Miranda, 

and Yaracuy; Map 209). La Divisoria de la Concepción; 1,300–

1,500 m.

Ornithidium Salisb. ex R. Br.

Ornithidium Salisb. ex R. Br., in W. T. Aiton, Hortus Kew., 2nd ed., 5: 210. 

1813. Maxillaria sect. Ornithidium (Salisb. ex R. Br.) Christenson, 

Richardiana 2(2): 52. 2002.

Siagonanthus Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 40. 1836. Maxillaria sect. 

Siagonanthus (Poepp. & Endl.) Christenson, Proc. 16th World Orchid 

Conf. 286. 2002.

Neourbania Fawc. & Rendle, J. Bot. 47: 125. 1909 (“Neo-urbania”).

Maxillaria sect. Reflexae Christenson, Proc. 16th World Orchid Conf. 285. 

2002.

Laricorchis Szlach., in Szlachetko and Śmiszek, Richardiana 7(1): 27. 2007 

[2006].

Primarily epiphytic but also lithophytic or terrestrial 

herbs; highly variable vegetatively. Monopodial, sympodial or 

sympodial as juveniles with monopodial adult shoots. Roots 

often orange-brown. Rhizomes creeping or attached only at 

base, producing erect shoots, rooting at internodes just below 

pseudobulbs or both. Pseudobulbs heteroblastic, ovoid or suborbicular, not or slightly compressed laterally, surface usually 

shiny and minutely reticulate (reminiscent of old varnish; best 

seen in living material), 1- or 2(–4)-foliate apically, usually enveloped by 2–5 imbricate sheaths, of which 1–3 internal ones 

have foliar blades, or pseudobulbs completely absent (e.g., Ornithidium serrulatum Lindl.). Leaves conduplicate, articulate; 

leaf blades often oblong-elliptic, occasionally linear or suborbicular, coriaceous to subfleshy. Inflorescences 1-flowered, 

usually fasciculate, rarely solitary, emerging from the sheaths 

subtending pseudobulbs or from sheaths of rhizomes, or stems 

erect; peduncles usually much longer than the pedicels; floral 

bracts inconspicuous, much shorter than the pedicels. Flowers small to (rarely) medium-sized, resupinate, usually brightly 

colored, yellow, red, orange, less commonly white or greenish, perianth segments fleshy, fibers absent, usually porrect, 

forming a tubular flower, subglobose or campanulate, often 

producing nectar at the base of the labellum; petals and sepals 

subequal, free, ovate or broadly elliptic, less commonly lanceolate, obtuse or broadly acute (rarely acute or acuminate), 

usually many-nerved. Labellum fleshier than other perianth 

segments, rigidly fused to the column foot or somewhat articulate, often geniculate, simple or 3-lobed, usually with a 

simple, conspicuous callus, fleshy in the middle of the disk; 

column cylindrical, straight or somewhat geniculate, with a 

short foot; anther terminal, operculate, incumbent, glabrous; 

pollinia 4, in 2 unequal pairs, usually white; stigmatic surface 

ventral, variable. Ovary terete or 3-angled, often much longer 

than peduncle. Capsules ellipsoid; dehiscent apically. [Epidendroideae: Cymbidieae: Maxillariinae.]

A genus of 55–60 species found in Mexico, Central America, the West Indies, and South America (Colombia, Venezuela, 

Trinidad and Tobago, the Guianas, Ecuador, Peru, Bolivia, and 

Brazil). The greatest species diversity occurs in the Andes; ~20 

species are found in Venezuela.

References. Atwood (1993); Blanco et al. (2007); 

Christenson (2002b); Pridgeon et al. (2009); Senghas (1993b, 

2002); Whitten et al. (2007).

Many species of Ornithidium with brightly colored, cuplike 

flowers fit the hummingbird-pollination syndrome, including the 

two species found in the park.

Authorship of the generic name Ornithidium occasionally 

is attributed to Salisbury (1812: 293), but he failed to provide 

a diagnostic character, stating only that the labellum of Ornithidium is more similar to that of Dendrobium Sw. than that of 

Cymbidium Sw. (see also Mabberley, 1980: 601).

KEY TO THE SPECIES OF ORNITHIDIUM

1a. Plants with large pseudobulbs at the base of long leafy stems (often missing in herbarium specimens), 5.5–9 cm long; leaf 

blades 7–10 × 0.1–1.8 cm; flowers in fascicles of 4–10, bright yellow with a red dot on the labellum; ovary 3–4 cm long, 

about 4× longer than the sepals; petals 1.7–2 mm wide, spathulate, much narrower than the sepals . . . . . O. miniatum

MAP 209. Oncidium abortivum occurrence in Venezuela.
NUMBER 100 • 181

1b. Plants with small pseudobulbs scattered over long rhizomes, 2–4 cm long; leaf blades 3–5 × 0.6–1 cm; flowers originating 

from the axils of the sheaths of the newly forming pseudobulbs, bright red-scarlet with purple at the apex of labellum 

and bright yellow at the base; ovary 1.5–2 cm long, ~2× longer than the sepals; petals 5–6 mm wide, broadly elliptic or 

ovate-elliptic, as wide or wider than the sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. ruberrimum

Ornithidium miniatum Lindl., Edwards’s Bot. Reg. 31: Misc. 

62. 1845. Maxillaria miniata (Lindl.) L. O. Williams, Caldasia 1(3): 14. 1941; Dunsterville and Garay, Venez. Orchids 

Ill. 2: 208–209. 1961; Foldats, Fl. Venez. 15(4): 486–488, 

fig. 741. 1970; Ortega et al., BioLlania 5: 48. 1987; Romero 

and Carnevali, Orchids Venez., 2nd ed., 2: 562. 2000; Dorr 

et al., Contr. U.S. Natl. Herb. 40: 51. 2000 [2001]; Fernández, Orquídeas Nat. Táchira 147. 2003; Morillo, in Morillo 

et al., Bot. Ecol. Monocot. Páramos Venez. 1: 236–238, fig. 

87, foto 44. 2011.

Maxillaria sp. A; Dorr et al., Contr. U.S. Natl. Herb. 40: 51. 2000 [2001].

Epiphytic, lithophytic or subterrestrial herbs; pseudobulbs 

to 9 cm long. Flowers axillary, in fascicles of 3–10. Sepals 6.5–10 

mm long. Petals bright yellow with a red dot on the apex of the 

labellum.

Found in South America (Colombia and Venezuela). In Venezuela, this species occurs in the Andes (Lara, Mérida, Táchira, 

and Trujillo), the Sierra de Perijá (Zulia), and the Cordillera de 

la Costa (Aragua, Distrito Federal, Falcón, Miranda, and Yaracuy; Map 210). Cloud forest on both slopes of Guaramacal; 

1,800–2,600 m.

Flower color in this species is extremely variable, and local 

populations with entirely red, orange or yellow flowers are 

known. So far, the only color form collected in the Guaramacal 

area displays yellow petals and sepals with a bright red dot on 

the apex of the labellum.

Ornithidium ruberrimum (Lindl.) Rchb. f., Ann. Bot. Syst. 6(4): 

489. 1861 [1863]. Scaphyglottis ruberrima Lindl., Orchid. 

Linden. 22. 1846; Morillo, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 1: 2386–2397, fig. 882. 2011. 

Maxillaria ruberrima (Lindl.) Garay, Bot. Mus. Leafl. 21: 

260. 1967; Dunsterville and Garay, Venez. Orchids Ill. 

6: 266–267. 1976; Ortega et al., BioLlania 5: 48. 1987; 

Romero and Carnevali, Orchids Venez., 2nd ed., 2: 584. 

2000; Dorr et al., Contr. U.S. Natl. Herb. 40: 51. 2000 

[2001]; Fernández, Orquídeas Nat. Táchira 151. 2003.

Maxillaria aggregata auct., non (Kunth) Lindl.; Foldats, Fl. Venez. 15(4): 

398–400, fig. 707. 1970; Ortega et al., BioLlania 5: 48. 1987.

Epiphytic, lithophytic or subterrestrial herbs; pseudobulbs 

5–10 cm distant on rhizomes. Flowers axillary, in groups of 1–3. 

Sepals 8–10 mm. Petals bright red with a purple labellum.

Found in South America (Colombia, Venezuela, and Brazil). 

In Venezuela, restricted to the Cordillera de Mérida (Mérida and 

Trujillo; Map 211). In the park, found on road cuts in cloud forest on both slopes of Guaramacal; (1,500–)1,800–2,600 m.

Ornithidium vagans Schltr. (=Camaridium bracteatum

(Schltr.) Schltr.), described from Cauca, Colombia, was included 

in the synonymy of Maxillaria ruberrima by Dunsterville and 

Garay (1976: 266), but the former name applies to a concept of 

the Cordillera Occidental of Colombia that has longer, narrower 

leaves, narrower petals, and more acute sepals and that is better 

accommodated in the related genus Camaridium.

MAP 210. Ornithidium miniatum occurrence in Venezuela. MAP 211. Ornithidium ruberrimum occurrence in Venezuela.
182 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Otoglossum (Schltr.) Garay & Dunst.

Otoglossum (Schltr.) Garay & Dunst., in Dunsterville and Garay, Venez. 

Orchids Ill. 6: 41. 1976. Odontoglossum subgen. Otoglossum Schltr., 

Repert. Spec. Nov. Regni Veg. Beih. 27: 109. 1924.

Oncidium subsect. Serpentia Kraenzl., in Engler, Pflanzenr. IV, 50 (Heft 80): 

167. 1922. Oncidium sect. Serpentia (Kraenzl.) Garay, Taxon 19: 455. 

1970.

Brevilongium Christenson, Richardiana 6(1): 47. 2006.

Ecuadorella Dodson & G. A. Romero, in Harling and Persson, Fl. Ecuador 

87: 345. 2010.

Epiphytic, lithophytic or terrestrial herbs, 20–100 cm tall, 

erect. Rhizomes creeping, well developed, thick, appressed to 

substrate or ascendent. Pseudobulbs heteroblastic, distant on 

rhizomes, 1- or 2-foliate apically, mostly oblong or ovoid to 

subspherical, clothed by several imbricate sheaths, the 1–3 innermost of which may have foliar blades. Leaves conduplicate, 

articulate, flat or concave, erect or spreading, mostly oblong, 

apically rounded or obtuse, often emarginate, fleshy or thickly 

coriaceous, sessile or with a ± developed pseudopetiole. Inflorescences originating from the base of the pseudobulb or among the 

enveloping sheaths, racemose, (1–)2–15(–25)-flowered, erect, 

dense; peduncles straight, terete, remotely sheathed, 1–3(–5)× 

longer than leaves; rachis straight or ± fractiflex; floral bracts 

often conspicuous but always shorter than the pedicellate ovary, 

navicular, often papery. Flowers resupinate, showy, widely 

spreading; perianth segments yellow or yellow-brown with 

darker brown or orange-brown blotches; labellum usually bright 

yellow. Sepals subequal, free, mostly obovate or elliptic-obovate, 

bases sometimes ± clawed, apices broadly obtuse to rounded. 

Petals subequal to the sepals. Labellum deflexed, the base replicate and adnate to column base, constricted about or below 

its middle (pandurate), below the constriction produced into a 

pair of spreading or retrorse basal lobes; central lobe obovate to 

suborbicular, rounded to truncate; disk with a complex callosity 

composed of several lamellae and teeth. Column short, semiterete, ascendent, forming a small, obtuse mentum with the ovary, 

footless, apically with 2 conspicuous, dolabriform wings; anther 

dorsal or subventral, operculate, incumbent; pollinia 2, waxy, 

stipe subquadrate, viscidium small; clinandrium cucullate, entire or erose; rostellum transverse; stigma ventral, subquadrate. 

Ovary terete, pedicellate. Capsules triangular in cross section. 

[Epidendroideae: Cymbidieae: Oncidiinae.]

A genus of 5 to 7 species found in Central America (Costa 

Rica and Panama) and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, and Brazil). Four species occur in Venezuela.

References. Neubig et al. (2012); Pridgeon et al. 

(2009); Senghas (1997d); Williams et al. (2001).

Superficially species of Otoglossum resemble those of 

Odontoglossum, especially in floral morphology.

Otoglossum coronarium (Lindl.) Garay & Dunst., in Dunsterville and Garay, Venez. Orchids Ill. 6: 41. 1976. Odontoglossum coronarium Lindl., Fol. Orchid. 1: Odontoglossum 

21. 1852.

Odontoglossum brevifolium auct., non Lindl.; Foldats, Fl. Venez. 15(5): 

202–205, excluding fig. 837. 1970.

Epiphytic herbs. Leaf blades to 30 cm long, 1 on top of the 

pseudobulb. Petals and sepals 3–6 cm long, dark brown-red with 

wavy, often yellowish margins. Labellum pandurate, yellow with 

red-brown on the constriction; callus brown and yellow.

Found in South America (Colombia, Venezuela, Ecuador, 

and Bolivia). In Venezuela, known only from the Cordillera de 

Mérida (Trujillo; Map 212). In the park, collected once in cloud 

forest without precise locality or elevation.

Platystele Schltr.

Platystele Schltr., Repert. Spec. Nov. Regni Veg. 8: 565. 1910.

Epiphytic, lithophytic or terrestrial herbs; caespitose or 

creeping, erect, small to diminutive. Rhizomes almost absent 

to creeping and branching. Upright portion of stem 1-foliate, 

very short to relatively elongate, with 1(to several) sheath at 

base. Leaves conduplicate, articulate, erect to prostrate, ovate, 

broadly elliptic or obovate, fleshy to thinly coriaceous, subsessile to (more commonly) subpetiolate. Inflorescences arising 

from near stem apex with an annulus just below the abscission 

layer; racemes short- to long-stalked, shorter than to conspicuously exceeding the subtending leaf, usually filiform; rachis 

often zigzag, less commonly straight, rarely very short and inflorescences capitate-umbellate; floral bracts inconspicuous. Flowers relatively small to tiny, usually resupinate, opening in slow 

succession or rarely ± simultaneously; perianth segments free, 

MAP 212. Otoglossum coronarium occurrence in Venezuela.
NUMBER 100 • 183

widely spreading, variable in shape but often acuminate, usually ± translucent, tinged with purple or lilac to yellow, usually 

1-nerved, membranous. Sepals similar or dorsal sepal longer 

than the others. Petals similar to sepals but often narrower and 

with ciliolate margins. Labellum simple to rarely sub-3-lobed, 

often ovate to narrowly ovate or elliptic, articulate with the column foot, margins often erose or ciliolate; column short and 

wide, membranous, widened apically; anther apical, incumbent, 

operculate; pollinia 2, narrowly pyriform to subspherical, stigmatic surface 2-lobed, transverse, near the apex of the column. 

Ovary filiform-pedicellate, smooth to costate. Capsules ellipsoid, usually crowned by the persistent perianth. [Epidendroideae: Epidendreae: Pleurothallidinae.]

Found in Mexico, Central America, the Greater Antilles, 

and South America (Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, and Brazil). The genus includes ~70 species, of 

which 8 occur in Venezuela.

References. Luer (1990); Pridgeon and Chase 

(2001); Pridgeon et al. (2001b, 2005).

Species of Platystele are typically shade loving.

Molecular support for the monophyly of Platystele is weak 

(Pridgeon and Chase, 2001; Pridgeon et al., 2001b).

Platystele pisifera (Lindl.) Luer, Selbyana 3: 232. 1977. Pleurothallis pisifera Lindl., Fol. Orchid. 9: Pleurothallis 36. 1859.

Platystele hypsitera Luer & R. Escobar, Orquideología 16: 14, t. 1983.

Epiphytic herbs, diminutive; densely caespitose. Leaf blades 

narrowly elliptical-obovate, 1–4 cm long. Inflorescences racemes, 3–6 cm long. Flowers translucent yellow or pink. Dorsal 

sepal 5–6 mm long. Labellum purple.

Found in South America (Colombia, Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, found only in the Andes 

(Mérida, Portuguesa, Táchira, and Trujillo; Map 213). Collected 

on both slopes of Guaramacal, including near La Divisoria de la 

Concepción and in Sector El Cafenol; 1,700–2,400 m.

Pleurothallis R. Br.

Pleurothallis R. Br., in W. T. Aiton, Hort. Kew., 2nd ed., 5: 211. 1813.

Epiphytic, lithophytic or terrestial herbs; caespitose to 

repent; rhizomes short or creeping; stems not thickened into 

pseudobulbs, cylindrical, triangular or angular in cross-section 

to laterally compressed, apically 1-foliate, occasionally apically 

proliferous (i.e., stems arising in the apex of others); leaves 

articulate, conduplicate and more commonly plane (dorsoventrally flattened), occasionally terete or laterally compressed, 

usually ovate, elliptic or obovate, less commonly linear or suborbicular, bases sessile or pseudopetiolate, sometimes cordate 

or decurrent on the stem, apices acute to rounded or emarginate, glabrous, sometimes glaucous, usually coriaceous. Inflorescences 1-flowered to racemose, arising from the apex of 

the stem associated with an annulus in multi-flowered inflorescences (annulus not evident in 1-flowered inflorescences), pedunculate or sessile, subtended by one spathe or not, flowering 

successively or simultaneously. Flowers resupinate or not, tiny 

to medium-sized or less commonly large; floral bracts variable, 

usually small and inconspicuous; pedicels usually short, articulate with the ovary. Sepals variable in shape, position, color, 

and ornamentation, membranous to fleshy; lateral sepals completely fused (less commonly only partially fused) into a synsepal of variable dimensions; dorsal sepal essentially free, similar 

to or very different than synsepal. Petals usually small and narrower than sepals, always free, sub-parallel to the labellum or 

widely divergent. Labellum usually fleshier than other perianth 

segments, simple or less commonly 3 (–5)-lobed, often provided 

with a glenion, plane, concave or convex, labellar base variably articulate with the base or the column or rigidly adnate 

to same and inflexible; column semiterete, relatively short, unwinged; clinandrium usually entire or not projecting forward, 

basally robust with a pedestal-shaped foot or foot sometimes 

absent; anther apical or subapical; pollinia 2, naked or with 

rudimentary caudicles, with or without a removable viscidium, 

pyriform or spherical; stigma transverse, 1- or 2-lobed. Ovary 

3-valvate, smooth to verrucose or muricate, keeled or crested. 

Capsules ellipsoid or obclavate. [Epidendroideae: Epidendreae: 

Pleurothallidinae.]

A neotropical genus of ~1,000 species found in Mexico, 

Central America, the West Indies, and South America (Colombia, Venezuela, Trinidad and Tobago, the Guianas, Ecuador, 

Peru, Bolivia, Brazil, and Argentina). The genus is most diverse 

in the Andes; ~50–60 species occur in Venezuela.

References. Luer (1986d, 1999, 2002b); Pridgeon 

and Chase (2001); Pridgeon et al. (2001b, 2005).

MAP 213. Platystele pisifera occurrence in Venezuela.
184 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

KEY TO THE SPECIES OF PLEUROTHALLIS

1a. Leaves petiolate or subpetiolate; inflorescences nutant or arching, 3–8-flowered; flowers translucent with purple or deep 

violet nerves; dorsal sepal >10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. dorrii

1b. Leaves sessile; inflorescences rigidly erect, at least 10-flowered (usually many more); flowers yellow, white or ± translucent green; dorsal sepal <10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. xanthochlora

Pleurothallis dorrii Luer, Lindleyana 12: 45, fig. 8. 1997.

Pleurothallis strobilifera auct., non F. Lehm. & Kraenzl.; Dorr et al., Contr. 

U.S. Natl. Herb. 40: 53. 2000 [2001].

Epiphytic herbs, erect, 20–36 cm tall. Leaf blades ellipticalobovate, 10–18 × 0.35–0.4 cm. Inflorescences nutant or arching, 

3–8-flowered, 5–11 cm long, much shorter than the subtending 

leaves. Flowers not resupinate, translucent with purple or deep 

violet nerves. Dorsal sepal 13–15 mm long.

Endemic to the Cordillera de Mérida of Venezuela (Portuguesa and Trujillo; Map 214). Found in La Divisoria de la Concepción and on the north slope of Guaramacal; 1,700–2,700 m.

This species was described from the nearby Montañas de 

Misisí (Trujillo state). It had been confused with Pleurothallis 

strobilifera F. Lehm. & Kraenzl., a species restricted to Bolivia.

Pleurothallis xanthochlora Rchb. f., Linnaea 22: 823. 1849; 

Dunsterville and Garay, Venez. Orchids Ill. 2: 312–313. 

1961; Foldats, Fl. Venez. 15(2): 449–450, fig. 359. 1970; 

Romero and Carnevali, Orchids Venez., 2nd ed., 3: 868. 

2000.

Pleurothallis revoluta auct., non (Ruiz & Pav.) Garay; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 53. 2000 [2001].

Epiphytic or subterrestrial herbs, 35–75 cm tall. Leaf blades 

oblong-elliptic to ovate, 10–22 × 4.5–7 cm. Inflorescences rigidly 

erect, 10-flowered (usually many more), 14–23 cm long. Flowers 

not resupinate, yellow, white or ± translucent green. Dorsal sepal 

4.5–5.5 mm long.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). In Venezuela, known from the Andes (Lara, 

Mérida, Portuguesa, and Trujillo) and the Cordillera de la Costa 

(Miranda; Map 215). In the park, found on both slopes of 

Guaramacal; 1,800–2,400 m.

Pleurothallopsis Porto & Brade

Pleurothallopsis Porto & Brade, Arq. Inst. Biol. Veg. 3: 133. 1937. Octomeria subgen. Pleurothallopsis (Porto & Brade) Luer, Monogr. Syst. Bot. 

Missouri Bot. Gard. 39: 80. 1991.

Restrepiopsis Luer, Selbyana 2: 199. 1978.

Epiphytic, lithophytic or terrestrial herbs; caespitose, erect 

to repent, usually inconspicuous. Rhizomes abbreviated. Stems 

1-leaved apically, erect, terete, enveloped by several scarious, 

tubular sheaths that are usually glabrous. Leaves conduplicate, 

articulate, linear-elliptic to suborbicular, sometimes attenuated 

into a pseudopetiole, coriaceous to fleshy coriaceous. Inflorescences originating from near the apex of the stem without an 

annulus, 1-flowered, solitary or in fascicles; peduncles short, terete; floral bracts tubular; pedicels short or rarely very elongate, 

MAP 214. Pleurothallis dorrii occurrence in Venezuela. MAP 215. Pleurothallis xanthochlora occurrence in Venezuela.
NUMBER 100 • 185

always longer than the floral bracts, articulate with the ovary. 

Flowers resupinate, usually greenish, yellowish or hyaline, often 

with reddish or purplish nerves. Perianth segments membranous, usually widely spreading. Sepals subequal or the dorsal 

sepal broader, free or more often the lateral sepals connate in 

the basal 1⁄2. Petals subequal to the sepals or more commonly 

smaller, frequently subparallel to the column. Labellum articulate to the column foot, erect, fleshier than the other perianth 

segments, subsimple or 3-lobed below the middle; lateral lobes 

much smaller than the central lobe, usually ± falcate, porrect 

and erect, ± embracing the column; central lobe usually flat or 

± concave, ovate-oblong, oblong to obovate; disk with 2 or 4 

longitudinal keels; column erect or ± curved, subterete, elongate; 

anther apical, incumbent, operculate, 2- or imperfectly 4-locular; 

pollinia 4, waxy, pyriform or ± clavate, viscidium very small; 

clinandrium shallow, mostly entire; rostellum transverse; stigma 

ventral, small. Ovary terete, often ridged. Capsules ellipsoid. 

[Epidendroideae: Epidendreae: Pleurothallidinae.]

Found in Central America and South America (Colombia, 

Venezuela, Ecuador, Peru, Bolivia, and Brazil). The genus includes ~15 species, 5 or 6 of which occur in Venezuela.

References. Luer (1991); Pridgeon and Chase 

(2001); Pridgeon et al. (2001b, 2005).

Pridgeon and Chase (2001: 255) argue that the molecular 

evidence for a sister relationship between Restrepiopsis Luer 

(four pollinia) and Pleurothallopsis (eight pollinia) does not 

support treating them as separate genera and that the proposed 

relationship with Octomeria R. Br. is strongly refuted. They further argue that the use of pollinia number alone is an unreliable 

generic character.

KEY TO THE SPECIES OF PLEUROTHALLOPSIS

1a. Labellum with well-developed basal lateral lobes; leaf blades linear-oblong or linear-elliptic, 4–10× longer than wide, 

equal to or longer than the stem, fleshy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. striata

1b. Labellum without well-developed lateral lobes, but erect basal margins; leaf blades lanceolate, 3–4× longer than wide, 

shorter than the stem, coriaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. tubulosa

Pleurothallopsis striata (Luer & R. Escobar) Pridgeon & M. W. 

Chase, Lindleyana 16: 255. 2001. Restrepiopsis striata Luer 

& R. Escobar, Orquideología 16: 43. 1983; Romero and 

Carnevali, Orchids Venez., 2nd ed., 3: 933. 2000; Dorr et 

al., Contr. U.S. Natl. Herb. 40: 53. 2000 [2001].

Restrepiella tubulosa auct., non (Lindl.) Garay & Dunst.; Foldats, Fl. Venez. 

15(2): 28–30, fig. 195. 1970; Ortega et al., BioLlania 5: 49. 1987.

Epiphytic herbs, 5–15 cm tall, erect. Leaf blades linear oblong or linear-elliptic, 4–7.5 × 0.5–0.7 cm. Flowers translucent 

yellow-green with purple veins in the dorsal sepal. Dorsal sepal 

6–8 mm long. Labellum yellow with purple veins.

Found in South America (Colombia and Venezuela). In Venezuela, confined to the Andes (Mérida, Táchira, and Trujillo; 

Map 216). In the park, found in cloud forest along the Boconó–

Guaramacal road; 1,850 m.

Pleurothallopsis tubulosa (Lindl.) Pridgeon & M. W. Chase, 

Lindleyana 16: 255. 2001. Pleurothallis tubulosa Lindl., 

Fol. Orchid. 9: Pleurothallis 19. 1859. Restrepiella tubulosa

(Lindl.) Garay & Dunst., in Dunsterville and Garay, Venez. 

Orchids Ill. 4: 266. 1966. Restrepiopsis tubulosa (Lindl.) 

Luer, Selbyana 2: 200. 1978; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 934. 2000; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 53. 2000 [2001]; Carnevali and RamírezMorillo, in Berry et al., Fl. Venez. Guayana 7: 547, fig. 456. 

2003; Fernández, Orquídeas Nat. Táchira 210. 2003; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 

1: 272–273, fig. 108, foto 55. 2011.

Pleurothallis viridula Lindl., Fol. Orchid. 9: Pleurothallis 19. 1859. Restrepiella viridula (Lindl.) Garay & Dunst., in Dunsterville and Garay, Venez. 

Orchids Ill. 4: 268. 1966; Foldats, Fl. Venez. 15(2): 30–32, fig. 196. 

1970. Restrepiopsis viridula (Lindl.) Luer, Selbyana 2: 200. 1978.

Epiphytic herbs, 6.5–21 cm tall, erect. Leaf blades lanceolate, 2–6 × 1–3 cm. Flowers greenish-yellow, often with 3 purple 

stripes in the dorsal sepal. Dorsal sepal 6–10 mm long; lateral 

sepals free or connate only at the very base. Labellum with a pair 

of small lateral lobes in the basal 1⁄4.

 Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, and Ecuador). In Venezuela, recorded from Amazonas, Aragua, Barinas, Bolívar, Distrito 

MAP 216. Pleurothallopsis striata occurrence in Venezuela.
186 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Federal, Mérida, Miranda, Táchira, Trujillo, and Zulia (Map 

217). Found above Boconó in cloud forest; 2,400 m.

We accept this species as part of the flora on the basis of 

a voucher cited by Dunsterville and Garay (1966: 268), which, 

given the locality and elevation, suggests their specimen might 

have been collected in Guaramacal National Park. We have, 

however, not seen their specimen, nor have we recollected the 

species within the boundaries of the park.

Pleurothallis tubulosa and P. viridula were published simultaneously. Luer (1991: 116) was the first to consider these two 

species synonyms, and he adopted the name Restrepiopsis tubulosa (≡Pleurothallopsis tubulosa). His choice of epithet must be 

followed (McNeill et al., 2012: Art. 11.5).

Ponthieva R. Br.

Ponthieva R. Br., in W. T. Aiton, Hort. Kew., 2nd ed., 5: 199. 1813.

Ocampoa A. Rich. & Galeotti, Ann. Sci. Nat., Bot., sér. 3, 3: 31. 1845.

Exalaria Garay & G. A. Romero, Harvard Pap. Bot. 4: 479. 1999.

Terrestrial or rarely epiphytic herbs; caespitose, erect, sympodial, each sympodium consisting of a rosette terminating in an 

inflorescence, rosettes dying annually after flowering is renewed 

by a new rosette that emerges from the base of the previous year. 

Roots few, fleshy, pubescent, emerging from stem base. Stems relatively short, completely covered by leaf sheaths. Leaves 2–7, not 

articulate, ovate to lanceolate, acute or obtuse, suberect, arching 

to prostrate on the substrate, often puberulent to pubescent, membranous or herbaceous; petiolate or subpetiolate; petioles often 

puberulent to pubescent. Inflorescences terminal, erect, few- to 

many-flowered racemes, peduncle and rachis usually puberulent 

to pubescent; peduncles terete, with 2–5 internodes, internodes 

subtended by remote tubular, appressed or more rarely subfoliaceous sheaths; rachis usually shorter than peduncle, terete, flowers spiraled. Floral bracts subfoliaceous, usually much shorter 

than the pedicellate ovaries, rarely subequal. Flowers not resupinate, small to medium, usually inconspicuous, delicate-textured, 

usually white or greenish-white, sometimes tinged with pink, lilac 

or brown in perianth segments and sepal apices or nerves brown 

or greenish; petals and labellum sometimes yellow. Sepals usually 

concave, adaxially puberulent, free, membranous or lateral sepals 

partially or completely fused into a broadly ovate or suborbicular 

synsepal; dorsal sepal usually thinner than the lateral ones. Petals adnate to the column above the receptacle, usually thinner 

than the sepals, oblique and basally attenuated in a ± conspicuous 

claw, connivent with the dorsal sepal and often constituting the 

most conspicuous part of the flower. Labellum fleshier and much 

smaller than the other segments of the perianth, adnate to the 

column above the receptacle in a complex structure, frequently 

concave and 3-lobed, central lobe usually shorter than the lateral 

ones or reduced to a mucro-like structure; column fleshy, massive, 

abbreviated in apical 1⁄2, lacking a foot; anther erect or subdorsal, articulate with the apex of the column, 2-locular, incumbent; 

pollinia 2, puberulent with a small, soft, triangular hamulus and 

a small, suborbicular viscidium; clinandrium short; rostellum 

erect, digitate, elongate, soft, fleshy, canaliculate. Ovary terete, 

pedicellate, usually much longer than the sepals, puberulent or 

pubescent. Capsules ovoid, ellipsoid or fusiform. [Orchidoideae: 

Cranichideae: Cranichidinae.]

A tropical and subtropical American genus of 25–30 species 

found in North America (USA), Mexico, Central America, the 

West Indies, and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Paraguay, and Argentina). Nine species 

occur in Venezuela.

References. Pridgeon et al. (2003); Salazar Chávez 

et al. (2009).

Salazar Chávez et al. (2009), in a molecular phylogenetic 

study, determined that Ponthieva was polyphyletic. They recognized that a clade containing the type of the genus would be 

monophyletic if the monotypic genera Ocampoa A. Rich. & 

Galeotti and Exalaria Garay & G. A. Romero were placed in 

synonymy, which they did, and this is the concept of the genus 

Ponthieva accepted here. Salazar Chávez et al. (2009) also noted 

that a second clade with Ponthieva species appeared to be allied to Baskervilla Lindl., which occurs in Central America and 

South America.

Ponthieva fertilis (F. Lehm. & Kraenzl.) Salazar, Ann. Bot. (Oxford), n.s., 104: 416. 2009. Goodyera fertilis F. Lehm. & 

Kraenzl., Bot. Jahrb. Syst. 26: 498. 1899. Cranichis fertilis

(F. Lehm. & Kraenzl.) Schltr., Repert. Spec. Nov. Regni Veg. 

Beih. 8: 115. 1921; Dunsterville and Garay, Venez. Orchids 

Ill. 5: 60–61. 1972; Dorr et al., Contr. U.S. Natl. Herb. 40: 

49. 2000 [2001].

Ophrys parviflora C. Presl, Reliq. Haenk. 1: 92. 1830 [1827], non Ponthieva 

parviflora Ames & C. Schweinf., Bot. Mus. Leafl. 4: 39. 1936. Exalaria 

MAP 217. Pleurothallopsis tubulosa occurrence in Venezuela.
NUMBER 100 • 187

parviflora (C. Presl) Garay & G. A. Romero, Harvard Pap. Bot. 4: 480. 

1999; Romero and Carnevali, Orchids Venez., 2nd ed., 1: 333. 2000; 

Fernández, Orquídeas Nat. Táchira 101. 2003; Morillo, in Morillo et 

al., Bot. Ecol. Monocot. Páramos Venez. 1: 176–178, fig. 532. 2011.

Cranichis pycnantha Schltr., Repert. Spec. Nov. Regni Veg. Beih. 7: 62. 1920; 

Foldats, Fl. Venez. 15(1): 419–421, fig. 157. 1969.

Terrestrial herbs, 20–45 cm tall. Leaf blades elliptic to 

oblong-lanceolate, 3–7 cm long. Flowers campanulate, white. 

Ovary 8–9 mm tall. Sepals 2–3.5 mm long.

Found in South America (Colombia, Venezuela, Ecuador, and Peru). In Venezuela, known from the Andes (Mérida, 

Táchira, and Trujillo) and the Cordillera de la Costa (Aragua, 

Distrito Federal, Falcón, and Miranda; Map 218). In the park, 

collected in forest on the south slope between Qda. Honda and 

El Santuario; 1,800–2,100 m.

Within Ponthieva, our species is somewhat anomalous by 

virtue of its small floral size, campanulate flowers, non-oblique 

petals, very short column with a large, completely dorsal anther, and segments of the perianth emerging from the base of 

the receptacle. These characters first caused this species to be 

described in Cranichis and later led to the creation of the monotypic Exalaria, but molecular phylogenetic analyses now support 

the placement of this species in Ponthieva.

Prescottia Lindl.

Prescottia Lindl., in Hooker, Exot. Fl. 2: t. 115. 1824 (“Prescotia”), nom. 

& orth. cons.

Terrestrial or rarely epiphytic herbs; acaulescent to shortly 

caulescent; glabrous to pubescent. Roots fleshy, often tuberous, 

fasciculate, usually villose. Leaves rosulate or less commonly 

somewhat cauline, frequently only a single leaf present, usually 

erect, less commonly the leaf appressed to the substrate; leaf 

blades widely elliptic or suborbicular to linear, prefoliation convolute, herbaceous, membranous to subfleshy, petiolate or sessile; petioles canaliculate. Inflorescences terminal, always erect 

and exceeding the leaves, simple, spicate with flowers appressed 

to the rachis; peduncles straight and ± rigid, often thick, variously 

covered with bract-like sheaths; rachis straight, densely manyflowered. Floral bracts ± equal in length to the ovary and pedicel. Flowers inconspicuous, not resupinate, short-lived, opening 

successively, erect, greenish to reddish. Sepals basally connate, 

forming a short cup, extended or more commonly reflexed, thintextured. Petals thinner than sepals, basally adnate to the sepaline cup, often circinate. Labellum fleshy, slipper-shaped, galeate 

or cochleate, often involving the column, basally with 2 retrorse 

auricles, claw adnate to sepaline cup; disk without a callus; column erect, short, fleshy, apically winged, basally extended into a 

short foot joining the labellar claw; anther erect, dorsal; pollinia 

4, without a tegula or caudicles; viscidium small; clinandrium 

erect, acuminate, margins connate with rostellum; rostellum 

laminar, smooth, emarginate, parallel to the anther and ± the 

same length; stigma entire, ventral. Ovary with thick, straight, 

fusiform to clavate pedicel, often longer than sepals. Capsules 

ovoid to ellipsoid. [Orchidoideae: Cranichideae: Cranichidinae.]

An American genus of ~25 species found in North America 

(USA), Mexico, Central America, the West Indies, and South 

America (Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina, and Uruguay). The genus is 

most diverse in Brazil; 6 species occur in Venezuela.

References. Álvarez-Molina and Cameron (2009); 

Pridgeon et al. (2003); Salazar Chávez et al. (2009).

According to a molecular phylogenetic analysis by Salazar 

Chávez et al. (2009), a monophyletic Prescottia can be achieved 

either by sinking the monotypic Pseudocranichis Garay in 

Prescottia or by removing Prescottia tubulosa (Lindl.) L. O. Williams from Prescottia. They chose to adopt the latter solution 

and applied the generic name Galeoglossum A. Rich. & Galeotti to the clade that includes Pseudocranichis and P. tubulosa. 

Galeoglossum, restricted to Mexico and Central America, can 

be distinguished from Prescottia by a labellum with incurved 

lateral margins that are open apically (not calceolate) and provided with a distinct apical lobule, the saddle-shaped stigma with 

two receptive areas separated by a central sterile area, and the 

hairpin-shaped, slender pollinia.

Prescottia stachyodes (Sw.) Lindl., Edwards’s Bot. Reg. 22: t. 

1915 [as “1916”]. 1836; Dunsterville and Garay, Venez. 

Orchids Ill. 1: 366–367. 1959; Foldats, Fl. Venez. 15(1): 

379–381, fig. 140. 1969; Romero and Carnevali, Orchids 

Venez., 2nd ed., 3: 891. 2000; Carnevali et al., in Berry et 

al., Fl. Venez. Guayana 7: 535–536. 2003. Cranichis stachyodes Sw., Prodr. 120. 1788.

Subterrestrial herbs, (25–)50–100 cm tall, very variable in 

size and leaf form. Petioles, peduncles, rachises, bracts, flowers, 

MAP 218. Ponthieva fertilis occurrence in Venezuela.
188 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

and fruit green, opaque red, brown, reddish or pink. Leaf blades 

elliptic-oblong to narrowly ovate or rounded-ovate, 5–20 × 2.5–

16 cm. Dorsal sepal 3–4 mm long.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Trinidad and Tobago, the 

Guianas, Ecuador, Peru, Bolivia, Brazil, and Argentina). In Venezuela, known from Amazonas, Anzoátegui, Aragua, Bolívar, Carabobo, Distrito Federal, Mérida, Miranda, Monagas, Portuguesa, 

and Yaracuy (Map 219). La Divisoria de la Concepción; 1,700 m.

This species is very variable throughout its extensive range, 

and we suspect that the taxon described here is actually part of 

a species complex.

Prosthechea Knowles & Westc.

Prosthechea Knowles & Westc., Fl. Cab. 2: 111. 1838.

Epithecia Knowles & Westc, Fl. Cab. 2: 167. 1839, nom. illeg.

Epidendrum subgen. Osmophytum Lindl., Edwards’s Bot. Reg. 25: Misc. 85. 

1839. Encyclia subgen. Osmophytum (Lindl.) Dressler & G. E. Pollard, 

Phytologia 21: 433. 1971. Encyclia sect. Osmophytum (Lindl.) Dressler 

& G. E. Pollard, Phytologia 21: 433. 1971.

Epidendrum subgen. Hormidium Lindl., J. Bot. (Hooker) 3: 81. 1841 [1840]. 

Hormidium (Lindl.) Heynh., Nom. Bot. Hort. 2: 880. 1841. Encyclia

sect. Hormidium (Lindl.) Dressler & G. E. Pollard, Phytologia 21: 433. 

1971. Prosthechea subgen. Hormidium (Lindl.) Charon & V. P. Castro, 

Richardiana 4(1): 29. 2003.

Anacheilium Rchb. ex Hoffmanns., Preiss-Verzeichn. Pflanzen 21. 1842.

Encyclia sect. Euchile Dressler & G. E. Pollard, Phytologia 21: 434. 1971. 

Euchile (Dressler & G. E. Pollard) Withner, Cattleyas & Relatives 5: 

137. 1998.

Pseudencyclia Chiron & V. P. Castro, Richardiana 4(1): 31. 2003.

Panarica Withner & P. A. Harding, Cattleyas & Relatives: Debatable Epidendrums 207. 2004.

Pollardia Withner & P. A. Harding, Cattleyas & Relatives: Debatable Epidendrums 217. 2004, nom. illeg.

Epiphytic, lithophytic or rarely terrestrial herbs; glycoside 

crystals present in all tissues of the majority of species. Roots relatively thick. Rhizomes short or creeping. Stems usually thickened 

into pseudobulbs (always in our species), heteroblastic (composed 

of a single thickened internode, sometimes 1 or more additional 

thickened internodes toward the base or apex), apically with 1−3

(–4) leaves. Leaves conduplicate, articulate, linear, elliptic or more 

commonly linear-oblong. Inflorescences terminal, 1-flowered (not 

our species), racemose or paniculate; peduncles subtended by a 

spathe or not, naked or partly covered by tubular sheaths. Flowers 

usually not resupinate, usually fragrant and long-lasting, showy to 

inconspicuous; floral bracts usually inconspicuous; perianth fleshy 

or fleshy-membranous, variously colored but more often greenish 

or yellow with brown or purple tints or white with the labellum 

with purple streaks and nerves. Sepals free, similar to one another. 

Petals free, usually similar to the sepals. Labellum fused to the 

column in the basal 1⁄3−1⁄2 portion, blade simple or 2-lobed; lateral lobes usually shorter and thinner than central lobe; disk of 

labellum often with callus, calli variable; column erect on pedicel, without columnar foot, parallel to the labellum; clinandrium 

3-toothed, teeth conspicuous, central tooth with an appendage; 

anther operculate, incumbent, 2-locular; pollinia 4, subreniform 

with caudicles of pollen material; rostellum transverse to the axis 

of the column; stigma ventral, semiorbicular, obcordate or transversely pandurate. Ovary pedicellate, smooth or variously ornamented, usually 3-winged. Capsules ellipsoid, fusiform, or ovoid, 

3-winged. [Epidendroideae: Epidendreae: Laeliinae.]

An American genus found in Mexico, Central America, the 

West Indies, and South America (Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, Brazil, and Paraguay). Prosthechea

is most diverse in Mexico and Brazil and moderately diverse in 

the Andes. There are ~100 species in the genus, 19 of which are 

found in Venezuela.

References. Berg and Chase (2004); Chiron 

(2003); Chiron and Castro Neto (2003); Higgins (1997, 1999); 

Higgins et al. (2003); Pridgeon et al. (2005).

Species of Prosthechea generally inhabit humid forest at low 

to intermediate elevations, and as a consequence, the genus is 

poorly represented in our flora.

When the genus Prosthechea first was proposed, it was 

considered to be monotypic and closely allied to the large genus 

Epidendrum. However, the name Prosthechea generally was not 

adopted until Higgins (1997, 1999) resurrected and expanded 

it, also transferring to Prosthechea species belonging to Encyclia

subgen. Osmophytum (Lindl.) Dressler & G. E. Pollard. Molecular evidence (Higgins et al., 2003) indicates that Prosthechea

is monophyletic, and various morphological characters also can 

be used to distinguish it from Encyclia s. str., including the fusiform, often flattened (vs. ovoid) pseudobulbs and the presence 

(vs. absence) of glycoside crystals.

MAP 219. Prescottia stachyodes occurrence in Venezuela.
NUMBER 100 • 189

KEY TO THE SPECIES OF PROSTHECHEA

1a. Pseudobulbs 9–18 cm long; sepals and petals yellow with brown or purplish-brown marks of various shapes; lateral 

lobes of labellum yellow or yellowish-cream, apices parallel to the central axis of the labellum, central lobe oblong; callus 

occupying most of the labellum disk and inserted at the base of the central lobe . . . . . . . . . . . . . . . . . . . P. brachychila

1b. Pseudobulbs 5–12 cm long; sepals and petals dark reddish-brown with yellowish or greenish apices; lobes of labellum 

white with red nerves, lateral lobes with retrorse apices, central lobe cordate; callus restricted to the basal 1⁄2 of the 

labellum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lindenii

Prosthechea brachychila (Lindl.) W. E. Higgins, Phytologia 82: 

376. 1997 [1998]; Romero and Carnevali, Orchids Venez., 

2nd ed., 3: 893. 2000; Fernández, Orquídeas Nat. Táchira 

202. 2003; Morillo, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 1: 263–264, fig. 102, foto 53. 2011. Epidendrum brachychilum Lindl., Orchid. Linden. 9. 1846; 

Dunsterville and Garay, Venez. Orchids Ill. 3: 86–87. 1965. 

Encyclia brachychila (Lindl.) Carnevali & I. Ramírez, Ernstia, ser. 1, 36: 9. 1986. Anacheilium brachychilum (Lindl.) 

Withner & P. A. Harding, Cattleya & Relatives: Debatable 

Epidendrums 52. 2004.

FIGURE 26D

Epidendrum hartwegii auct., non Lindl.; Foldats, Fl. Venez. 15(3): 268–271, 

fig. 477. 1970, pro parte.

Epiphytic herbs, 30–40 cm tall. Leaf blades oblong, 15–20 × 

1.3–1.8 cm. Inflorescences 15–17 cm tall, 8–10-flowered. Dorsal 

sepal 12–13 mm long.

Found in South America (Colombia and Venezuela). In Venezuela, confined to the Andes (Lara, Mérida, Táchira, and Trujillo; 

Map 220). Cloud forest on the north slope of the park, including 

cloud forest below the Páramo del Pumar; 1,850–2,600 m.

Prosthechea brachychila is similar and certainly related to P. 

hartwegii (Lindl.) W. E. Higgins, with which it is sympatric (at 

least in Venezuela). The latter species, however, has smaller flowers that are more opaque and less showy and a labellum with 

much wider lateral lobes. Prosthechea hartwegii occurs from 

Venezuela to Bolivia in the Andes, whereas P. brachychila is restricted to the Cordillera Oriental in Colombia and the Andes in 

Venezuela.

Prosthechea lindenii (Lindl.) W. E. Higgins, Phytologia 82: 379. 

1997 [1998]; Romero and Carnevali, Orchids Venez., 2nd 

ed., 3: 901. 2000; Fernández, Orquídeas Nat. Táchira 204. 

2003; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 265–266, fig. 103, foto 54. 2011. Epidendrum 

lindenii Lindl., Ann. Mag. Nat. Hist., ser. 1, 12: 397. 1843; 

Dunsterville and Garay, Venez. Orchids Ill. 2: 96, 132–133. 

1961; Foldats, Fl. Venez. 15(3): 306–308, fig. 492. 1970. 

Encyclia lindenii (Lindl.) Carnevali & I. Ramírez, Ernstia, 

ser. 1, 36: 9. 1986. Anacheilium lindenii (Lindl.) Hágsater, 

in Escobar, Nat. Colombian Orchids 1: 28. 1990.

Epiphytic or terrestrial herbs, 15–20 cm tall. Leaf blades 

oblong or oblong-lanceolate, 10–22 × 1–2.5 cm. Inflorescences 

10–40 cm tall, 4–18-flowered. Dorsal sepal 15–20 mm long.

Found in South America (Colombia and Venezuela). In Venezuela, found in the Andes (Lara, Mérida, Táchira, and Trujillo) 

MAP 220. Prosthechea brachychila occurrence in Venezuela. MAP 221. Prosthechea lindenii occurrence in Venezuela.
190 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

and in the Sierra de Perijá (Zulia; Map 221). Cloud forest on the 

north slope of the park and forest below the Páramo del Pumar; 

1,950–2,600 m.

Psilochilus Barb. Rodr.

Psilochilus Barb. Rodr., Gen. Spec. Orchid. 2: 272. 1882.

Terrestrial herbs; sympodial. Rhizomes terete, basally creeping and leafless, apically erect and spirally leaved, rooting at 

the internodes, usually branching below the middle after anthesis. Roots few, fleshy, pubescent. Leaves distichous, sheathing, 

petiolate or clasping, not articulate, prefoliation convolute, 2–5 

per stem, distant and leaving most of the stem naked, elliptic, 

ovate or lanceolate, green concolorous or variegated with silvery 

longitudinal zones, plicate, membranous. Inflorescences terminal, racemose (rarely branching), 2–10(–20)-flowered, usually 

shorter than the leaves; floral bracts conspicuous, navicular or 

subfoliaceous. Flowers resupinate, subcampanulate, short-lived, 

successive or rarely several flowering simultaneously; perianth 

segments free, membranous, whitish or greenish, often with purple lines or tinges; petals and sepals narrowly lanceolate to narrowly obovate, subequal, equal or lateral sepals oblique, dorsally 

carinate. Petals usually narrower than the sepals, often falcate. 

Labellum completely free from the column, equal to petals; blade 

narrowed toward the base, wider and sharply 3-lobed above the 

middle, less commonly lobed below the middle, central lobe sessile or with a conspicuous claw; disk with several longitudinal 

keels; column elongate, terete, tapering toward the base, usually 

arched; anther terminal, erect, articulate with the apex of the 

column, 2-locular; pollinia 4, granulose; stigma ventral. Ovary 

pedicellate, terete or subtrigonous, arched. Capsules ellipsoid to 

obovoid, often ± trigonous. [Epidendroideae: Triphoreae.]

A genus of 9 species found in Mexico, Central America, the 

West Indies, and South America (Colombia, Venezuela, Guyana, Ecuador, Peru, Bolivia, and Brazil). Four species occur in 

Venezuela.

References. Pridgeon et al. (2005); Rothacker 

(2007).

Species of Psilochilus are generally shade loving.

Psilochilus macrophyllus (Lindl.) Ames, Orchidaceae 7: 45. 

1922; Foldats, Fl. Venez. 15(1): 159–161, fig. 56. 1969, pro 

parte. Pogonia macrophylla Lindl., Ann. Mag. Nat. Hist., 

ser. 3, 1: 335. 1858.

Terrestrial herbs, to 40 cm tall. Leaves sheathing, sessile, 

clasping or lower leaves sometimes short-petiolate; leaf blades 

ovate, 5–10 × 2.5–7 cm. Inflorescences erect, 1- to severalflowered. Flowers successive. Sepals equal, ~11–12 × 2 mm, 

greenish yellow or green. Petals linear falcate, equal to sepals. 

Labellum cuneate, ~12–15 mm long, 3-lobed.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Bolivia, and Brazil). In 

Venezuela, known only from the Andes (Trujillo; Map 222). Our 

collections are from Qda. Segovia on the north slope of the park; 

2,100–2,550 m.

Psilochilus macrophyllus is often confused with P. physurifolius (Rchb. f.) Løjtnant, and in Venezuela the latter generally 

has been considered a synonym of the former. The two species, 

however, can be distinguished by vegetative characters; our species has amplexicaul or clasping leaves (some lower leaves are 

occasionally short-petiolate), whereas P. physurifolius has only 

petiolate leaves. Both species are found in the Venezuelan Andes. 

It appears that material from the Venezuelan Guayana referred 

to a broadly circumscribed P. macrophyllus (see Foldats, 1969; 

Carnevali and Ramírez-Morillo in Berry et al., 2003; Carnevali 

et al. in Hokche et al., 2008) is P. physurifolius, which was described from Guyana. We have not reviewed the material from 

the Cordillera de la Costa assigned by these same authors to P. 

macrophyllus, and we do not know its correct identity.

Pterichis Lindl.

Pterichis Lindl., Gen. Sp. Orchid. Pl. 444. 1840.

Terrestrial herbs; caespitose. Rhizomes short or absent. 

Roots tuberous, linear-fusiform, fasciculate at the base of the 

rosette. Plants rosulate, 1 or 2(3) leaves per rosette, leaf blades 

herbaceous, attenuate basally into a canaliculate pseudopetiole, 

often withered and absent at anthesis (i.e., plants hysteranthous). 

Inflorescences lateral; peduncles erect, rigid, simple, pubescent, 

with several sheaths; sheaths tubular, remote and leaving most of 

the peduncle naked; rachis in the apical 1⁄4 of the inflorescence, 

lax to densely spiciform. Floral bracts conspicuous, naviculiform, 

usually not exceeding the pedicellate ovary in length. Flowers not 

MAP 222. Psilochilus macrophyllus occurrence in Venezuela.
NUMBER 100 • 191

resupinate, successive or 2–4 in simultaneous anthesis, lasting 

1 or 2 days; perianth segments subfleshy, pubescent externally, 

usually reddish, brown or yellow with red or red-brown streaks. 

Sepals free or lateral sepals somewhat connate basally, similar or 

the lateral sepals somewhat wider, usually elliptic or lanceolate. 

Petals free, narrower than sepals, straight to somewhat falcate. 

Labellum free to base, fleshier than the other perianth segments, 

deeply concave (in living material), simple or with an apical 

lobe, broadly transverse-ovate in general outline, often with 

verrucose-glandular (papillate) structures along margins, apical 

lobe usually recurved, labellar margins partially clasping the column; disk variously pubescent or glandular-pubescent; column 

short and lacking a foot; anther dorsal, erect; clinandrium small; 

rostellum horizontal; pollinia 4, ovoid-clavate, brittle, with small 

terminal caudicles attached to a small hamulus; viscidium terminal, relatively small; stigma ventral, entire. Ovary pedicel thickened, pubescent, equal to or conspicuously longer than its bract. 

Capsules oblong or ellipsoid. [Orchidoideae: Cranichideae: 

Cranichidinae.]

A neotropical genus of 15–20 species found in Central 

America, the Greater Antilles, and South America (Colombia, 

Venezuela, Ecuador, Peru, Bolivia, and Argentina). The greatest 

species diversity is in the tropical Andes. Five species are known 

from Venezuela.

References. Morales (1986); Pridgeon et al. (2003); 

Salazar Chávez et al. (2009).

Species of Pterichis are sun-loving plants found in open 

areas at high elevations (2,500–4,000 m).

Limited molecular sampling suggests that Pterichis is monophyletic (Salazar Chávez et al., 2009). Salazar Chávez et al. 

(2009) also speculate that the morphologically similar monotypic Fuertesiella Schltr., restricted to the Greater Antilles, might 

prove to be synonymous with Pterichis. A revision of this relatively small genus could resolve this question.

Pterichis multiflora (Lindl.) Schltr., Bot. Jahrb. Syst. 45: 389. 

1911; Dunsterville and Garay, Venez. Orchids Ill. 4: 256–

257. 1966; Foldats, Fl. Venez. 15(1): 399–401, fig. 149. 

1969. Acraea multiflora Lindl., Orch. Linden. 26. 1846.

Pterichis diuris auct., non Rchb. f.; Dunsterville and Garay, Venez. Orchids 

Ill. 6: 40. 1976; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 

914. 2000.

Terrestrial herbs, 16–55 cm tall. Leaf blades oblong or 

oblong-lanceolate, 4–15 × 1.1–2.9 cm. Floral bracts 12–13 × 6.5 

mm. Flowers yellow-green with deep-brown or reddish streaks. 

Sepals 3-nerved, pubescent; dorsal sepal 9–11 × 3.4–3.6 mm; lateral sepals 8–9 × 3.9–4.1 mm. Petals 9–11 × 1.9–2.1 mm (5× longer than wide), 3-nerved. Labellum 6.5–7 × 10–11 mm, densely 

pubescent internally.

Found in South America (Colombia, Venezuela, and Ecuador). In Venezuela, known only from the Andes (Mérida, 

Táchira, and Trujillo; Map 223). In the park, found in páramo 

near the Laguna del Pumar; ~3,000 m.

Scaphyglottis Poepp. & Endl.

Scaphyglottis Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 58. 1835 [1836], nom. 

cons.

Hexisea Lindl., J. Bot. (Hooker) 1: 7. 1834, nom. rej.

Cladobium Lindl., Intr. Nat. Syst. Bot., 2nd ed., 446. 1836.

Hexadesmia Brongn., Ann. Sci. Nat., Bot., sér. 2, 17: 44. 1842.

Tetragamestus Rchb. f., Bonplandia (Hannover) 2: 21. 1854.

Reichenbachanthus Barb. Rodr., Gen. Sp. Orchid. 2: 164. 1881 [1882] 

(“Reichembachanthus”).

Epiphytic, rarely lithophytic or subterrestrial, herbs; small 

and inconspicuous to large; usually caespitose; erect to arching 

or subpendulous. Roots relatively thin, white, numerous, arising 

from the base of pseudobulbs. Rhizomes short. Pseudobulbs heteroblastic or partially homoblastic, basally surrounded by few to 

several sheaths, of which only rarely 1 or 2 have foliar blades, 

usually narrowly cylindrical to narrowly cylindrical-fusiform, 

sessile to stipitate, 1- or 2(3)-leaved apically, new pseudobulbs 

originating either from the bases of old pseudobulbs or their apices, often proliferous, making plants architecturally complex as 

they consist of 2–6 superimposed levels of pseudobulbs. Leaves 

conduplicate, articulate, linear, oblong, oblong-elliptic to ovateelliptic, acute to obtuse, usually unequally 2-lobed and mucronate apically, leaf blades concolorous or mottled with purple 

or reddish, sessile or rarely subpetiolate. Inflorescences terminal, originating from the apices of the most recent pseudobulbs 

and/or the lowermost pseudobulbs when superimposed, usually 

1-flowered, rarely racemose or paniculate, commonly reduced, 

when solitary, frequently fasciculate and successive and the 

flowers arising from a mass of sheaths and bracts. Floral bracts 

MAP 223. Pterichis multiflora occurrence in Venezuela.
192 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

usually inconspicuous. Flowers resupinate or not, usually inconspicuous, sometimes relatively attractive, subcampanulate to 

widely open, relatively short-lived, often cleistogamous; perianth 

segments membranous to ± fleshy, white, green, yellow, pink 

or bright orange, often mottled or with red or purple nerves, 

less commonly the entire flower or labellum violet-blue. Sepals 

subsimilar or sides of lateral sides slightly wider and somewhat 

oblique, partially fused to the sides or foot of the column, forming a mentum; dorsal sepal free. Petals similar to sepals but usually shorter and narrower, rarely wider. Labellum articulate or 

rigidly attached to the apex of the column foot, erect, ± spreading or recurved, variously sessile or with a simple or lobed claw, 

often emarginate to 2-lobed apically; disk ± ecallose; labellar base 

fused with columnar base to form a nectariferous cup (in a group 

of species with orange flowers and ornithophilous pollination 

syndrome); column short to relatively elongate, erect or forming 

an angle with the pedicellate ovary, without a wing or variously 

winged in the upper 1⁄2, basally expressed as a ± well-developed 

columnar foot; anther terminal, operculate, incumbent; pollinia 

4, 6 or 8, waxy, laterally compressed, with caudicles, yellow or 

cream (in orange-flowered species); clinandrium entire or variously 3-lobed; rostellum transverse, stigma ventral. Ovary terete, pedicellate, often relatively elongate, sometimes recurved. 

Capsules ellipsoid to subglobose, often beaked. [Epidendroideae: 

Epidendreae: Laeliinae.]

A neotropical genus of ~60 species found in Mexico, Central America, the West Indies, and South America (Colombia, 

Venezuela, the Guianas, Ecuador, Peru, Bolivia, and Brazil). The 

genus is most diverse in Costa Rica and Panama, but 21 species 

are found in Venezuela.

References. Berg et al. (2009); Dressler et al. 

(2004); Pridgeon et al. (2005).

As a consequence of molecular phylogenetic analyses 

(Dressler et al., 2004), the circumscription of Scaphyglottis has 

been expanded, and the genus now includes species formerly 

assigned to Hexisea Lindl. (a generic name rejected in favor of 

Scaphyglottis), Hexadesmia Brongn., Reichenbachanthus Barb. 

Rodr., Cladobium Lindl., and Tetragamestus Rchb. f.

Scaphyglottis summersii L. O. Williams, Bot. Mus. Leafl. 9: 14, 

t. 3, figs. 4–7. 1940; Romero and Carnevali, Orchids Venez., 

2nd ed., 3: 962. 2000; Fernández, Orquídeas Nat. Táchira 

214. 2003.

Epiphytic herbs, to 45 cm long. Leaf blades linear or linearlanceolate, (2–)5–8 cm × 2–5 mm. Inflorescences 1-flowered, 

single or fascicled. Flowers pale purplish-brown. Dorsal sepal 

1–1.5 cm long.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). In Venezuela, known only from the Cordillera de Mérida (Trujillo; Map 224). Collected on the north 

slope of the park near the Laguna de Aguas Negras and on 

the south slope between El Campamento and Qda. Honda; 

1,900–2,100 m.

Schlimmia Planch. & Linden

Schlimmia Planch. & Linden, in Linden, Établ. Linden, Prix-Courant 7: 5. 

1852.

Primarily epiphytic or terrestrial herbs; erect; sympodial; 

caespitose. Roots few, relatively thick. Pseudobulbs ovoid or subcylindrical, erect, heteroblastic, 1(2)-foliate, enclosed by scarious 

sheaths that lack blades, smooth. Leaves plicate, glossy gray-green, 

elliptic, long-petiolate, prominently 5-nerved below. Inflorescences 

solitary, pendulous or seemingly creeping, 2–5(–10)-flowered; 

peduncles terete, filiform, covered with a few appressed, remote 

bracts. Flowers fleshy, not resupinate, white, lasting to 10 days 

(relatively long-lived in comparison to other Stanhopeinae). Floral bracts conspicuous. Dorsal sepal free, lateral sepals fused to 

form a helmet or deeply concave sac, ovate. Petals free, oblongspathulate, falcate, less fleshy than the sepals. Labellum shorter 

than the column, hidden in the sepaline sac, divided into an entire 

hypochile with rugose, yellow callus and smooth, elliptic-ovate 

epichile; column semiterete, auriculate on distal 1⁄2, with a conspicuous foot at the base to which the lateral sepals are attached; 

anther subventral, operculate, pollinaria composed of 2 yellow, 

oblong, dorsiventrally flattened pollinaria, long and slender stipe, 

and semilunate viscidium; rostellum long filiform; stigmatic surface narrow, transverse. Ovary pedicellate, longer than the sepals, 

6-furrowed. [Epidendroideae: Cymbidieae: Stanhopeinae.]

A neotropical genus of 6 or 7 species restricted to South 

America (Colombia, Venezuela, Guyana, Ecuador, Peru, and Bolivia). A single species occurs in Venezuela.

References. Gerlach (1999a); Pridgeon et al. 

(2009); Sweet (1973); Whitten et al. (2000).

MAP 224. Scaphyglottis summersii occurrence in Venezuela.
NUMBER 100 • 193

Schlimmia alpina Rchb. f. & Warsz., Bonplandia (Hannover) 2: 

98. 1854; Romero and Carnevali, Orchids Venez., 2nd ed., 

3: 967. 2000 (“Schlimia”); Llamozas et al., Libro Rojo Fl. 

Venez. 459. 2003 (“Schlimia”).

Schlimmia trifida Rchb. f., Gard. Chron., n.s., 2: 708. 1876 (“Schlimia”); 

Dunsterville and Garay, Venez. Orchids Ill. 3: 282–283. 1965 

(“Schlimia”); Foldats, Fl. Venez. 15(4): 110–112. 1970 (“Schlimia”).

Epiphytic or subterrestrial herbs; pseudobulbs narrowly 

pyriform. Leaf blades elliptic, 15–20 cm long. Inflorescences 

10–15 cm long. Flowers white. Dorsal sepal 2–2.5 cm long.

Found in South America (Colombia, Venezuela, Guyana, 

and Ecuador). In Venezuela, known from the Andes (Lara and 

Trujillo) and the Cordillera de la Costa (Yaracuy; Map 225).

Inclusion of this species in the flora is based on a sight record made in the park by G. Gerlach (pers. comm.).

Sobralia Ruiz & Pav.

Sobralia Ruiz & Pav., Fl. Peruv. Prodr. 120. 1794, nom. cons. prop.

Terrestrial, lithophytic or epiphytic herbs; relatively small 

to large and robust; caespitose; when epiphytic occasionally subpendulous, ascending or patent. Rhizomes short, thick. Stems 

terete or laterally compressed, sometimes slightly woody at base, 

usually simple but sometimes somewhat branched or proliferous, becoming scandent, partly or completely enclosed by leaf 

sheaths. Leaves distichous, plicate, articulate, sessile on the 

sheaths, elliptic or ovate-elliptic, rarely linear, ± rigid to subcoriaceous, less commonly thin; sheaths tubular and clasping the stem 

or funnel-shaped and greatly inflated, glabrous, verruculose, rugose or variously pubescent. Inflorescences terminal or axillary, 

sessile or pedunculate, racemose or paniculate; peduncles terete, 

laterally flattened or angled. Floral bracts cucullate, inconspicuous to large and subfoliaceous. Flowers showy, medium-sized to 

very large, resupinate, short-lived (lasting 1 or 2 [3] days), usually white, yellow, lilac, pink or purple, less commonly greenish 

or brown, opening widely or subcampanulate; sepals and petals 

similar or petals slightly wider than sepals, free, membranous 

or somewhat subfleshy. Labellum entire or more rarely lobed, 

usually more conspicuous than other perianth segments, basally 

surrounding the column, apically opening in a blade ornamented 

with bristles, keels or dentate crests; column without a foot, relatively large and narrow, with membranous, apical wings that terminate in a pair of falcate, dentate or digitate structures on both 

sides of the stigma; anther terminal, incumbent, long, operculate, 

2-locular; pollinia 8, 4 in each locule, dusty-grained in texture, 

rarely somewhat waxy; stigmatic surface reniform, transverse; 

rostellum large, convex. Capsules relatively large-oblongoid or 

linear-oblongoid. [Epidendroideae: Sobralieae.]

As traditionally construed, a genus of ~100 species found 

in Mexico, Central America, and South America (Colombia, 

Venezuela, the Guianas, Ecuador, Peru, Bolivia, and Brazil). The 

genus is most diverse in Central America and the Andes; ~20 

species occur in Venezuela.

References. Dressler (2002); Neubig et al. (2011); 

Pridgeon et al. (2005).

A molecular phylogenetic study of the Sobralieae (Neubig 

et al., 2011) identified a core Sobralia clade that was monophyletic, but several species of Sobralia sect. Sobralia fell outside 

this clade. This is problematic nomenclaturally as the type of 

the genus is among these outliers, and this is the basis for a conservation proposal by Dressler et al. (2011), which has received 

preliminary approval (Applequist, 2012: 1113).

The one species encountered in Guaramacal falls within the 

core Sobralia clade.

Sobralia violacea Linden ex Lindl., Orchid. Linden. 26. 1846; 

Dunsterville and Garay, Venez. Orchids Ill. 1: 400–401. 

1959; Foldats, Fl. Venez. 15(1): 197–199, fig. 73B. 1969; 

Romero and Carnevali, Orchids Venez., 2nd ed., 3: 993. 

2000; Carnevali and Ramírez-Morillo, in Berry et al., Fl. 

Venez. Guayana 7: 574. 2003.

Sobralia yauaperyensis auct., non Barb. Rodr.; Dunsterville and Garay, 

Venez. Orchids Ill. 2: 324–325. 1961; Foldats, Fl. Venez. 15(1): 199–

201, fig. 73A. 1969.

Terrestrial or rarely epiphytic herbs; stems erect, to 1.5(–2) 

m tall. Leaf blades lanceolate to oblong-lanceolate, 15–25 × 4–7 

cm. Inflorescences terminal. Flowers 1(–3) successive, emerging 

from a mass of sheaths; perianth segments usually pink or pale 

lilac-violet, more rarely dark violet or totally white. Sepals 5–7 

cm long. Labellum tube and calli yellow.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, Bolivia, and Brazil). Widespread in Venezuela, including 

the Andes (Barinas, Lara, Mérida, Portuguesa, Táchira, and 

MAP 225. Schlimmia alpina occurrence in Venezuela.
194 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Trujillo), the Sierra de Perijá (Zulia), the Cordillera de la Costa 

(Falcón and Miranda), and the Venezuelan Guayana (Amazonas, 

Bolívar, and Delta Amacuro; Map 226). In the park, found on 

the north slope below Páramo del Pumar and on the south slope 

between El Campamento and Qda. Honda; 1,900–2,100 m.

This species is sun loving, locally common, and often found 

on road cuts.

Specklinia Lindl.

Specklinia Lindl., Gen. Sp. Orchid. Pl. 8. 1830. Pleurothallis subgen. Specklinia (Lindl.) Garay, Orquideología 9: 121. 1974.

Epiphytic, lithophytic or terrestrial herbs. Rhizomes short, 

creeping. Stems erect, usually very short, terete or ventrally canaliculate, often covered by 1 to several appressed, imbricate tubular sheaths. Leaves erect with respect to the stem, commonly 

elliptic to linear or obovate, rarely subcylindric, bases usually attenuate, apices minutely 3-toothed, coriaceous, fleshy or fleshycoriaceous. Inflorescences terminal, racemose, solitary or (less 

commonly) in fascicles, shorter than the subtending leaves to 

several times longer, erect or less commonly patent, associated 

with an annulus and emerging from its apex and just below the 

leaf articulation, flowers opening successively or simultaneously. 

Floral bracts membranous, usually reduced. Flowers membranous (compared to most Pleurothallidinae), commonly resupinate, variously colored, small or medium-sized. Sepals almost 

free or lateral sepals fused for varying lengths (often completely 

fused) into a plane, or more commonly, concave synsepal supporting the labellum, externally smooth or verruculose, dorsal 

sepal often more slender than synsepal and differently shaped, 

apices often caudate. Petals less fleshy and conspicuously smaller 

than sepals, variously shaped, parallel to the column or somewhat divergent, margins entire or often dentate to fimbriate or 

laciniate. Labellum articulate with the column foot and rarely 

with a pair of tiny lobes or retrorse teeth at the base of each side 

of the articulation with the column foot, blade deeply 3-lobed 

to more commonly simple, lateral lobes relatively small, erect, 

apical lobe entire, verruculose or pubescent, margins entire to 

more commonly erose to laciniate, disk often with 2 or more 

longitudinal calli in the basal or middle portion, often concave; 

column elongated, semicylindrical and ventrally concave, erect 

or ± arched, foot well developed; clinandrium usually erose or 

dentate to fimbriate; anther operculate, subventral; stigma and 

rostellum ventral; pollinia 2. Ovary articulate with pedicel, terete 

or subtriangular; pedicel usually filiform and much longer and 

thinner than the ovary. Capsules globose to obovoid. [Epidendroideae: Epidendreae: Pleurothallidinae.]

A neotropical genus of ~200 species found in Mexico, Central America, the West Indies, and South America (Colombia, 

Venezuela, Ecuador, Peru, Bolivia, Brazil, and Argentina). The 

genus is most diverse in the Andes; 5 or more species are found 

in Venezuela.

References. Luer (1986d, 2006); Pridgeon and 

Chase (2001); Pridgeon et al. (2005).

Specklinia lacks obvious morphological synapomorphies, 

but the majority of species are small caespitose plants with short 

stems with an annulus and a hinged labellum. The limits of the 

genus and its relationship to other pleurothallids are still a subject of debate (Luer, 2004, 2006; Pridgeon et al., 2005). Neither of the two species found in Guaramacal would remain in 

Specklinia as defined by Luer (2004, 2006), who dismembered 

the genus, dividing it into 23 vegetatively similar genera.

KEY TO THE SPECIES OF SPECKLINIA

1a. Inflorescences including peduncle, rachis and floral bracts laterally compressed; flowers relatively large with sepals >1 cm 

long; sepals green, orange or red, 3–5-nerved; dorsal sepal elliptic, at least 6 mm wide midlength, much longer than the 

petals; petals emerging from the basal portion of the column; labellum deeply geniculate in living material, shorter than 

or equal to column in length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. endotrachys

1b. Inflorescences including peduncle and rachis terete, floral bracts concave; flowers relatively small; sepals translucent or 

creamy, 1-nerved; dorsal sepal linear, <1 mm wide midlength, slightly longer than the petals; petals emerging from the 

apex of the floral receptacle; labellum flat in living material, longer than column in length . . . . . . . . . . . . S. zephyrina

MAP 226. Sobralia violacea occurrence in Venezuela.
NUMBER 100 • 195

Specklinia endotrachys (Rchb. f.) Pridgeon & M. W. Chase, 

Lindleyana 16: 257. 2001. Pleurothallis endotrachys Rchb. 

f., Linnaea 41: 95. 1877 [1876]; Dunsterville and Garay, 

Venez. Orchids Ill. 3: 250–251. 1965; Foldats, Fl. Venez. 

15(2): 265–266, fig. 279. 1970; Ortega et al., BioLlania 

5: 49. 1987; Romero and Carnevali, Orchids Venez., 2nd 

ed., 3: 779. 2000; Dorr et al., Contr. U.S. Natl. Herb. 40: 

52. 2000 [2001]. Empusella endotrachys (Rchb. f.) Luer, 

Monogr. Syst. Bot. Missouri Bot. Gard. 95: 258. 2004.

Epiphytic herbs, to 50 cm tall (including the inflorescences). 

Leaf blades oblanceolate to narrowly oblanceolate, 12–20 cm × 

1–3 mm. Inflorescences 10–45 cm long, straight or curved, fewflowered. Sepals 2–2.5 cm long.

Found in Mexico, Central America, and South America 

(Colombia and Venezuela). In Venezuela, known only from the 

Cordillera de Mérida (Barinas and Trujillo; Map 227). In the 

park, found along the Boconó–Guaramacal road; ~2,450 m.

Specklinia zephyrina (Rchb. f.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95: 265. 2004. Pleurothallis zephyrina

Rchb. f., Bonplandia (Hannover) 3: 71. 1855; Romero and 

Carnevali, Orchids Venez., 2nd ed., 3: 870. 2000; Dorr et 

al., Contr. U.S. Natl. Herb. 40: 52. 2000 [2001]; Carnevali 

and Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 7: 

523. 2003; Fernández, Orquídeas Nat. Táchira 197. 2003. 

Trichosalpinx zephyrina (Rchb. f.) Luer, Phytologia 54: 

398. 1983. Muscarella zephyrina (Rchb. f.) Luer, Monogr. 

Syst. Bot. Missouri Bot. Gard. 105: 124. 2006.

Pleurothallis setigera Lindl., Fol. Orchid. 9: Pleurothallis 38. 1859; Dunsterville and Garay, Venez. Orchids Ill. 4: 242–243. 1966; Foldats, Fl. Venez. 

15(2): 412–414, fig. 342B. 1970; Ortega et al., BioLlania 5: 49. 1987.

Tiny epiphytic herbs, to 5 cm tall (including inflorescences). Leaf blades narrowly obovate, 1–2.5(–3.5) × 2.5–

4.5 cm. Inflorescences 5–10 cm long, suberect. Dorsal sepal 

7–8.5 mm long.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). In Venezuela, known only from the Andes 

(Lara, Mérida, Táchira, and Trujillo; Map 228). In the park, 

found along the Boconó–Guaramacal road; ~2,550 m.

Stelis Sw.

Stelis Sw., J. Bot. (Schrader) 1799(2): 239. 1800, nom. cons.

Apatostelis Garay, Bot. Mus. Leafl. 27: 185. 1979 [1980] (“Apatoselis”).

Epiphytic, terrestrial or lithophytic herbs; caespitose or 

creeping, minute to fairly large (in comparison to other Pleurothallidinae). Rhizomes prostrate or ascendent, sometimes distinctly pendent, often branching and then plants mat-forming. 

Stems almost absent to very elongated, shorter than to much 

longer than the single apical leaf, terete, covered partially or 

totally by sheaths, in many species apically proliferous (and 

then producing chains of superposed stems). Leaves solitary on 

top of the stems, articulate, usually elliptic or elliptic-obovate, 

sometimes linear or oblong, thinly coriaceous to fleshy or even 

subterete, sessile to conspicuously subpetiolate. Inflorescences 

racemes, few- to many-flowered, solitary to several, flowers 

opening successively or simultaneously, originating on or just 

below apex of the stem from a swollen annulus, basally often 

subtended by a spathe, rachis straight or zigzag, rarely very abbreviated and then inflorescences umbellate. Flowers usually 

resupinate, spreading widely to campanulate, triangular sepals 

variously connate, forming a triangular-shaped flower made of 

MAP 227. Specklinia endotrachys occurrence in Venezuela. MAP 228. Specklinia zephyrina occurrence in Venezuela.
196 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

3 triangles fused at base, the central portion of the triangle occupied by an apparatus composed of the column, the 2 petals, 

and the labellum; perianth segments membranous to thinly coriaceous, usually white, green, yellow or purple, often variously 

pubescent. Sepals connate basally, 1-, 3-, 5-, 7- or 9-veined, 

similar or the lateral sepals fused together to form a bilabiate 

flower, or the dorsal sepal larger than the other 2. Petals usually 

much smaller than the sepals, fleshy, transverse, margin thickened, fitting snugly in the shallow tube formed by the base of 

the sepals and closely flanking the labellum and column. Labellum small, thick, usually shaped like 1⁄4 of a sphere, the uppermost plane of the labellum resting beneath the column and often 

bearing a glenion and 1 or few keels or ridges, basally sessile 

on the column. Column short and broad, dilated apically; anther apical, incumbent, operculate; pollinia 2, usually pyriform; 

stigma apical, transverse, 1- or 2-lobed, when 2-lobed, the lateral lobes protrude to either side of the anther under the flap on 

the rostellum, stigma lobes often so elongated that the receptive 

surfaces are held far to either side of the anther. Capsules ellipsoidal, fusiform or cylindrical. [Epidendroideae: Epidendreae: 

Pleurothallidinae.]

A neotropical genus of ~900 species found in Mexico, Central America, the West Indies, and South America (Colombia, 

Venezuela, the Guianas, Ecuador, Peru, Bolivia, and Brazil). The 

genus is most diverse in the Andes and southern Central America; ~80 species occur in Venezuela.

References. Duque (2003a, 2003b, 2008); Garay 

(1979); Karremans et al. (2013); Pridgeon and Chase (2001); 

Pridgeon et al. (2001b, 2005).

The flowers of Stelis species often are sensitive to light or 

temperature and open or close at particular times of the day in 

response to the appropriate stimulus.

The generic concept presented here is that of Stelis s. str., 

which corresponds to Clade F in the molecular phylogeny of 

Karremans et al. (2013). As such, the species are united by 

terminal, racemose, fascicled, few- to many-flowered inflorescences; triangular, almost radially symmetrical flowers with ± 

identical sepals; diversely connate sepals that are much larger 

than the petals and labellum; greatly reduced petals with thick 

margins; a thickened labellum similar to the petals; and a very 

short, unwinged column with an apical stigma and anther (Pridgeon et al., 2001b; Karremans et al., 2013). An alternative 

molecular classification (Pridgeon and Chase, 2001; Pridgeon 

et al., 2001b) accepts Stelis s.l. and includes Crocodeilanthe in 

synonymy. Luer (2004, 2006) rejected this latter interpretation 

and, using mostly morphological characters, argued for an even 

greater splitting of the genus than is accepted in this flora or by 

Karremans et al. (2013).

KEY TO THE SPECIES OF STELIS

1a. Rhizomes elongate, plants creeping or ascending . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Plants diminutive; leaf blades ≤0.6 cm wide; sepals <2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pusilla

2b. Plants larger; leaf blades ≥0.6 cm wide; sepals >2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Inflorescences solitary, 2 or 3× longer than the leaves in length; stems clustered and ascending, rhizomes growing 

upward; labellum semicircular (as seen from above), as long as wide; sepals pubescent, subequal . . . S. ascendens

 3b. Inflorescences 1(–3), shorter than, subequal to or slightly exceeding the leaves in length; rhizomes clearly creeping; labellum subquadrate-obovate (as seen from above), longer than wide; sepals glabrous; dorsal sepal larger 

than lateral sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. atra

1b. Rhizomes short, plants caespitose, never creeping or ascending . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4a. Lateral sepals more fused to each other than to the dorsal sepal; synsepal concave . . . . . . . . . . . . . . . . . . . . . . . . 5

 5a. Petals and labellum terminating in a long acumen; dorsal sepal 1.7–2.5 mm long . . . . . . . . . . . . . S. biserrula

 5b. Petals and labellum not terminating in a long acumen; dorsal sepal 4–6 mm long . . . . . . . . . . . . . . S. vulcani

4b. Lateral sepals as fused to each other as they are to the dorsal sepal; synsepal flat or convex . . . . . . . . . . . . . . . . . 6

 6a. Plants very large, 25–30(–70) cm tall; leaf blades ≥5 cm wide; sepals 5-nerved, very wide (broadly deltoid) and 

almost completely fused, sinuses of synsepal very shallow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lindenii

 6b. Plants smaller, to 15(–35) cm tall; leaf blades ≤5 cm wide; sepals 3-nerved, narrower (lanceolate or ovate) and 

fused <1⁄2 their length, sinuses of synsepal very deep . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 7a. Sepals revolute so that the flower is convex; petals apiculate . . . . . . . . . . . . . . . . . . . . . . . . . . S. hylophila

 7b. Sepals not revolute so that the synsepal is flat or concave; petals not apiculate . . . . . . . . . . . . . . . . . . . . 8

 8a. Mature plants producing 4–8 inflorescences per stem; stem conspicuously longer (2 or 3×) than the leaves 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. striolata

 8b. Mature plants producing 1–3 inflorescences per stem; stem either shorter or longer than the leaves, but 

never 2 or 3× longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

 9a. Inflorescences very lax; rachis internodes much longer than the sepals; sepals lanceolate, ~2× longer 

than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

 10a. Stems >5 cm long; usually multiple inflorescences per stem; labellum 3-lobed, longer than wide 

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. chamaestelis

 10b. Stems 1–3 cm long; inflorescences 1(–2) per stem; labellum simple, as wide as long. . . . . S. nitens
NUMBER 100 • 197

 9b. Inflorescences denser; rachis internodes shorter than to ± equal to the sepals; sepals broadly ovate, 

as long as wide or at most 1.5× longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

 11a. Stems very short, 1–2 cm long in adult plants; inflorescences shorter than or exceeding the 

leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. humilis

 11b. Stems longer, at least 3 cm long in adult plants; inflorescences always exceeding the leaves . . . 12

 12a. Inflorescences dense; floral bracts imbricate; sepals unequal . . . . . . . . . . . . . S. oblonga

 12b. Inflorescences lax; floral bracts contiguous or remote; sepals subequal . . . . . . . . . . . 13

 13a. Inflorescences emerging from a spathe appressed to the pseudopetiole; labellum with 

an apiculus; petal margins glandular-verruculose . . . . . . . . . . . . . . . . S. argentata

 13b. Inflorescences emerging from a spathe diverging from the pseudopetiole; labellum 

without an apiculus; petal margins entire or only cellular-verruculose . . . . . . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. atroviolacea

Stelis argentata Lindl., Edwards’s Bot. Reg. 28: Misc. 64. 1842; 

Dunsterville and Garay, Venez. Orchids Ill. 1: 414–415. 

1959; Foldats, Fl. Venez. 15(2): 62–64, fig. 203. 1970; 

Romero and Carnevali, Orchids Venez., 2nd ed., 3: 1005. 

2000; Carnevali and Ramírez-Morillo, in Berry et al., Fl. 

Venez. Guayana 7: 584. 2003; Fernández, Orquídeas Nat. 

Táchira 220. 2003.

Epiphytic herbs, to 30 cm tall. Leaf blades oblong to oblongoblanceolate, (2–)6.5–12.5 × 0.7–2.5(–4) cm. Inflorescences solitary, erect, 10–25 cm long, usually greatly exceeding the leaves, 

many-flowered. Flowers greenish-yellow to reddish-brown. Sepals patent, subequal, broadly elliptic ovate, 2–5.5 × 2–5.5 mm, 

± pubescent on margins and villous-papillose on lamina. Petals 

broadly cuneate-rhomboid. Labellum fleshy, globular.

Found in Central America and South America (Colombia, 

Venezuela, the Guianas, Ecuador, Peru, Bolivia, and Brazil). 

Widespread in Venezuela (Amazonas, Anzoátegui, Apure, Aragua, Bolívar, Delta Amacuro, Distrito Federal, Mérida, Monagas, 

Sucre, Táchira, and Trujillo; Map 229). In the park, found along 

the Boconó–Guaramacal road; 2,100–2,300 m.

This species as currently construed is very variable morphologically and has a very wide range.

Stelis ascendens Lindl., in Hooker, Companion Bot. Mag. 2: 

353. 1836; Foldats, Fl. Venez. 15(2): 64–67, fig. 204A. 1970 

(“ascendants”); Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 281, fig. 112. 2011.

Epiphytic herbs, to 20 cm tall. Leaf blades oblong-elliptic to 

narrowly oblong-obovate, 2.7–7.5 × 0.7–1.6 cm. Inflorescences 

solitary, ± erect, 8–22 cm long, greatly exceeding the leaves, secund, many-flowered. Flowers green with purple or violet. Sepals 

± patent, ovate, subequal, 2–2.6 × 2–2.6 mm, pilose, margins 

sometimes appearing ciliate. Petals narrowly elliptic. Labellum 

fleshy, semicircular.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, occurs in the Andes (Mérida, Táchira, 

MAP 229. Stelis argentata occurrence in Venezuela. MAP 230. Stelis ascendens occurrence in Venezuela.
198 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

and Trujillo) and in the Cordillera de la Costa (Sucre; Map 230). 

In the park, collected in cloud forest on both slopes of Guaramacal; 1,700–2,700 m.

Whether or not this is the correct name for this species needs 

to be resolved as Duque (2008) considers Stelis ascendens, described from Peru, to be a synonym of S. jamesonii Lindl. (J. Bot. 

(Hooker) 1: 11. 1834), which he restricts to Colombia, Ecuador, 

and Peru.

Stelis atra Lindl., Orchid. Linden. 3. 1846; Garay and Dunsterville, Venez. Orchids Ill. 4: 290–291. 1966; Foldats, Fl. 

Venez. 15(2): 66–68, fig. 204B. 1970; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 1006. 2000; Morillo, in 

Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 282–

283, fig. 113. 2011. Stelis hallii var. atra (Lindl.) Garay & 

C. Schweinf., Fieldiana, Bot., n.s., 33: 11. 1970; Dorr et al., 

Contr. U.S. Natl. Herb. 40: 54. 2000 [2001].

Epiphytic or terrestrial herbs, <10 cm tall. Leaf blades lanceolate to oblanceolate, 4.5–8 × 0.9–2 cm. Inflorescences 1(–3), 

erect, 3.5–8 cm long, slightly shorter or longer than the leaves, 

many-flowered. Flowers purple, reddish-purple, wine-colored or 

pinkish. Sepals unequal, glabrous; dorsal sepal broadly ovate, 

2.5–3 × 2.5–3 mm; lateral sepals ovate, 1.7–2 × 1.5–2 mm. Petals 

trapezoid obtuse. Labellum subquadrate-obovate.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). In Venezuela, apparently restricted to the 

Cordillera de Mérida (Mérida, Portuguesa, and Trujillo; Map 

231). In the park, known from cloud forest on both slopes of 

Guaramacal; 1,700–3,100 m.

Stelis atroviolacea Rchb. f., Bonplandia (Hannover) 3: 70. 1855; 

Dunsterville and Garay, Venez. Orchids Ill. 2: 330–331. 

1961; Foldats, Fl. Venez. 15(2): 68–70, fig. 205A. 1970; 

Romero and Carnevali, Orchids Venez., 2nd ed., 3: 1007. 

2000.

Epiphytic herbs, 18–27 cm tall. Leaf blades ovate-oblong 

to spathulate, 7–12 × 1.6–2.5 cm. Inflorescences 1(or more?), 

to 18 cm long, shorter or slightly longer than the leaves, manyflowered. Flowers deep purple to clear green. Sepals subequal, 

elliptic-ovate, 2–3 × 2–3 mm, shortly villous. Petals reniform. 

Labellum oblong, concave.

Found in South America (Colombia, Venezuela, and Ecuador). In Venezuela, found in the Andes (Lara, Mérida, Táchira, 

and Trujillo) and in the Cordillera de la Costa (Aragua, Carabobo, and Miranda; Map 232). Guaramacal, without precise 

locality; 2,400–2,600 m.

Stelis biserrula Lindl., Fol. Orchid. 8: Stelis 16. 1858 [1859]; 

Dunsterville and Garay, Venez. Orchids Ill. 3: 298–299. 

1965; Foldats, Fl. Venez. 15(2): 73–76, fig. 207. 1970; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 

1: 283–284. 2011.

Stelis triseta auct., non Lindl.; Romero and Carnevali, Orchids Venez., 2nd 

ed., 3: 1053. 2000; Dorr et al., Contr. U.S. Natl. Herb. 40: 54. 2000 

[2001]; Fernández, Orquídeas Nat. Táchira 225. 2003.

Epiphytic or terrestrial herbs, to 16 cm tall; stems long trailing. 

Leaf blades lanceolate to narrowly elliptic, (3.5–)6–10 × 0.6–1.8

(–2.5) cm. Inflorescences 1(–3), erect, 5–10(–25) cm tall, exceeding 

the leaves or not, many-flowered. Flowers secund, nutant, green, 

MAP 231. Stelis atra occurrence in Venezuela. MAP 232. Stelis atroviolacea occurrence in Venezuela.
NUMBER 100 • 199

translucent green, pale green, pale yellow, whitish-green or whitish 

translucent. Sepals unequal, glabrous; dorsal sepal elliptic-ovate, 

1.7–2.5 × 1.6–3 mm, lateral sepals smaller, 1.6–4 × 1.6–3 mm. 

Petals rhombic. Labellum trapezoidal, apiculate.

Found in Central America and South America (Colombia, 

Venezuela, Ecuador, Peru, and Bolivia). Venezuelan records are 

all from the Andes (Mérida, Portuguesa, Táchira, and Trujillo; 

Map 233). In the park, found in cloud forest on both slopes of 

Guaramacal; 1,800–2,700 m.

We follow Luer (2009) in considering Stelis biserrula to be 

distinct from S. bicornis Lindl. (=S. triseta Lindl.), which nonetheless is a closely related species restricted to the Andes of Ecuador, Peru, and Bolivia. In fact, Luer (2009: 53) states that S. 

biserrula is “possibly only a variation of the common and widely 

distributed Stelis bicornis Lindl. with a small habit and flowers.”

Dunsterville and Garay (1965) illustrated Stelis biserrula

and vouchered their illustration with Dunsterville & Dunsterville 720 (AMES), a specimen collected between Boconó and 

Guaramacal. Ortega et al. (1987) and Dorr et al. (2000), however, cited Dunsterville & Dunsterville 772 (VEN) as a voucher 

for S. biserrula, which makes us now suspect that the label on the 

duplicate in Caracas was miscopied as that collection number 

(i.e., 772) is a paratype of Encyclia ivonae Carnevali & G. A. 

Romero, a species found in the Venezuelan Guayana.

Stelis chamaestelis (Rchb. f.) Garay & Dunst., in Dunsterville 

and Garay, Venez. Orchids Ill. 4: 292. 1966; Foldats, Fl. 

Venez. 15(2): 77–78, fig. 208B. 1970; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 1011. 2000; Morillo, in 

Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 284, 

fig. 114. 2011. Pleurothallis chamaestelis Rchb. f., Linnaea 

22: 825. 1849. Apatostelis chamaestelis (Rchb. f.) Garay, 

Bot. Mus. Leafl. 27: 187. 1979 [1980].

Epiphytic herbs, to 35 cm tall. Leaf blades narrowly ellipticoblong, 3.5–12 × 1.4–1.7 cm. Inflorescences 1 to several, fasciculate, ± erect, 7–12(–19) cm long, exceeding the leaves, lax, 

many-flowered. Flowers white or yellowish-green. Sepals equal, 

narrowly elliptical, 1.2–2 × 0.6–1.1 mm, glabrous. Petals obtusely 

rhombic. Labellum subquadrate, 3-lobed, longer than wide.

Found in South America (Colombia and Venezuela). In 

Venezuela, restricted to the Andes (Lara, Mérida, Portuguesa, 

Táchira, and Trujillo; Map 234). In the park, collected along 

the Boconó–Guaramacal road and on the south slope in Sector 

Paramito–La Aguadita and near La Divisoria de la Concepción; 

1,700–1,900(–3,000) m.

One Guaramacal collection (Cuello et al. 1604) describes 

the flowers as “moradas,” but that is certainly an error.

Stelis humilis Lindl., Fol. Orchid. 8: Stelis 10. 1858 [1859]; 

Garay and Dunsterville, Venez. Orchids Ill. 1: 420–421. 

1959; Foldats, Fl. Venez. 15(2): 104–106, fig. 220. 1970; 

Romero and Carnevali, Orchids Venez., 2nd ed., 3: 1023. 

2000; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 288–2893, fig. 117. 2011.

Epiphytic herbs, 6–20 cm tall. Leaf blades narrowly elliptic to oblong-elliptic, 3–12 × 0.3–1 cm. Inflorescences solitary, 

± erect, 4–12 cm long, shorter or longer than the leaves, densely 

many-flowered. Flowers nutant, greenish-brown. Sepals equal, 

ovate, 2–3.5 × 1.7–3 mm, ± hairy, especially near margins. Petals 

subrhomboid. Labellum subtrapezoidal to subquadrate.

MAP 233. Stelis biserrula occurrence in Venezuela. MAP 234. Stelis chamaestelis occurrence in Venezuela.
200 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Found in South America (Colombia and Venezuela). In Venezuela, found in the Andes (Lara, Mérida, Táchira, and Trujillo) 

and in the Cordillera de la Costa (Aragua, Cojedes, Distrito 

Federal, Falcón, Miranda, and Yaracuy; Map 235). In the park, 

found on the north slope in Qda. Segovia; ~1,850–1,900 m.

Stelis hylophila Rchb. f., Bonplandia (Hannover) 3: 241. 1855; 

Garay and Dunsterville, Venez. Orchids Ill. 4: 302–303. 

1966; Foldats, Fl. Venez. 15(2): 106–108, fig. 221. 1970; 

Romero and Carnevali, Orchids Venez., 2nd ed., 2: 1024. 

2000; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 290–291, fig. 118. 2011. Apatostelis hylophila (Rchb. f.) Garay, Bot. Mus. Leafl. 27: 189. 1979 [1980].

Epiphytic herbs, to 30 cm tall. Leaf blades oblong or narrowly elliptic-oblong, 6–12 × 0.8–2.5 cm. Inflorescences 1 to 

several, 5–11 cm long, equal to or exceeding leaves, densely 

many-flowered. Flowers translucent white with a dark center (or 

yellowish-green). Sepals equal, ovate-elliptic, 1.1–1.4 × 1–1.3 

mm, papillate at the apices, often reflexed. Petals cuneate, apiculate. Labellum subquadrate, rounded or subcuneate.

Found in South America (Colombia, Venezuela, Ecuador, and 

Peru). In Venezuela, evidently restricted to the Andes (Mérida, 

Táchira, and Trujillo; Map 236). Collected along the Boconó–

Guaramacal road on the north slope of Guaramacal; 2,100–2,200 m.

Stelis lindenii Lindl., Orchid Linden. 3. 1846 (“Lindeni”); Foldats, 

Fl. Venez. 15(2): 118–120, fig. 227. 1970; Morillo, in Morillo 

et al., Bot. Ecol. Monocot. Páramos Venez. 1: 194. 2011.

Stelis purdiaei auct., non Lindl.; Romero and Carnevali, Orchid Venez., 2nd 

ed., 3: 1042. 2000.

Epiphytic herbs, 25–35(–70) cm tall. Leaf blades oblong, 

10–20 × 3.5–7 cm. Inflorescences 1 to several, 15–25 cm long, 

as long as or exceeding the leaves, many-flowered. Flowers 

greenish-yellow suffused with pink. Sepals equal, broadly deltoid and almost completely fused, 4–6 × 3–4.5 mm, minutely 

pubescent or glabrous. Petals rounded to subquadrate. Labellum 

subquadrate to suboblong.

Found in South America (Colombia and Venezuela). In 

Venezuela, found only in the Andes (Lara, Mérida, Táchira, and 

Trujillo; Map 237). Infrequently collected in the park, where it 

has been found near the Boconó–Guaramacal road and along 

the track from El Campamento to Qda. Honda; 1,900–2,100 m.

This is the largest of the Stelis species found in Guaramacal.

MAP 236. Stelis hylophila occurrence in Venezuela.

MAP 235. Stelis humilis occurrence in Venezuela.

MAP 237. Stelis lindenii occurrence in Venezuela.
NUMBER 100 • 201

Stelis nitens Rchb. f., Bonplandia (Hannover) 2: 22. 1854; Garay 

and Dunsterville, Venez. Orchids Ill. 4: 310–311. 1966; Foldats, Fl. Venez. 15(2): 130–132, fig. 232. 1970; Romero and 

Carnevali, Orchids Venez., 2nd ed., 2: 1036. 2000; Fernández, Orquídeas Nat. Táchira 224. 2003; Morillo, in Morillo 

et al., Bot. Ecol. Monocot. Páramos Venez. 1: 295, fig. 122, 

foto 59. 2011.

Epiphytic herbs, to 25 cm tall. Leaf blades oblong-elliptic to 

± spathulate, 3.5–11 × 1–2 cm. Inflorescences 1(–2), erect, 10–18 

cm long, as long as or exceeding the leaves, ± lax, many-flowered 

apically. Flowers green or greenish-yellow. Sepals subequal, narrowly ovate-lanceolate, 2.5–3.5 × 1.8–2 mm. Petals cochleateoblong. Labellum subquadrate.

Found in South America (Colombia, Venezuela, and Ecuador). In Venezuela, principally found in the Andes (Mérida, 

Portuguesa, Táchira, and Trujillo), but the type is from Caracas (Distrito Federal), and there is a collection from Zulia (Map 

238). In the park, found in El Cafenol and Qda. Segovia on the 

north slope of Guaramacal; 1,800–2,300 m.

Stelis oblonga (Ruiz & Pav.) Willd., Sp. Pl. 4(1): 139. 1805; 

Dunsterville and Garay, Venez. Orchids Ill. 4: 312–313. 

1966; Foldats, Fl. Venez. 15(2): 134–136, fig. 234A. 1970; 

Romero and Carnevali, Orchids Venez., 2nd ed., 2: 1038. 

2000; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 296–297, fig. 123. 2011. Humboltia oblonga Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 1: 236. 1798 

(“Humboldtia”).

?Stelis alba Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 291. 1815 [1816]; 

ibid. [qu.] 1: 363. 1815 [1816]; Dunsterville and Garay, Venez. Orchids 

Ill. 5: 288. 1972; Romero and Carnevali, Orchids Venez., 2nd ed., 2: 

1004. 2000.

Epiphytic or lithophytic herbs, 7–22 cm tall. Leaves oblong 

to oblong-elliptic, 7–9 × 2–3.5 cm. Inflorescences 1 to several, 

erect, 6.5–17 cm long, many-flowered. Flowers congested, yellow, green or greenish-yellow. Sepals unequal, broadly ovate, 

papillose at the apices; dorsal sepal 1.9–2.5 × 1.7–2.2 mm; lateral sepals 1.6–2 × 1.6–2 mm. Petals subrhomboid. Labellum 

suborbicular to subrhomboid.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). Infrequently collected in the Venezuelan Andes (Lara, 

Mérida, Táchira, and Trujillo; Map 239). In the park, found near 

the Laguna de los Cedros and on both slopes of Guaramacal; 

1,850–2,350 m.

We suspect that a specimen (Dunsterville 653, AMESfragment) collected on the Boconó–Guaramacal road at 2,100–

2,400 m and used to illustrate Stelis alba (Dunsterville and 

Garay, 1972) is the same as our S. oblonga, but that needs to be 

confirmed. The two names are sometimes considered synonyms, 

but both species are recognized by Duque (2008), who although 

he acknowledges that their floral structures are similar, argues 

without elaboration that their habits differ.

Stelis pusilla Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 289. 

1815 [1816]; ibid. [qu.] 1: 361. 1815 [1816]; Foldats, Fl. 

Venez. 15(2): 145–147, fig. 238. 1970; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 1043. 2000; Carnevali 

and Ramírez-Morillo, in Berry et al., Fl. Venez. Guayana 7: 

586. 2003; Morillo, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 1: 300, fig. 126. 2011.

Epiphytic herbs, 6–10(–22) cm tall. Leaf blades linearoblanceolate to lanceolate, 1.5–4.5 × 0.2–0.6 cm. Inflorescences solitary, 2–6 cm long, erect, many-flowered. 

Flowers yellow-green. Sepals subequal, ovate to ovate-elliptic, 

MAP 238. Stelis nitens occurrence in Venezuela. MAP 239. Stelis oblonga occurrence in Venezuela.
202 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

1.3–1.8 × 1.2–1.8 mm. Petals cuneate, slightly apiculate. Labellum ± suborbicular.

Found in the Greater Antilles and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, and Brazil). In Venezuela, 

found in the Andes (Lara, Mérida, Táchira, and Trujillo), the 

Cordillera de la Costa (Miranda and Yaracuy), and the Venezuelan Guayana (Bolívar; Map 240). We have a single record from 

Qda. Jirajara where the specimen was collected on a log that had 

fallen across a cascading stream; 2,000 m.

Stelis striolata Lindl., Fol. Orchid. 8: Stelis 4. 1858 [1859]; Dunsterville and Garay, Venez. Orchids Ill. 4: 318–319. 1966; 

Foldats, Fl. Venez. 15(2): 156–158, fig. 244. 1970; Romero 

and Carnevali, Orchids Venez., 2nd ed., 3: 1051. 2000; Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 

1: 303–304, fig. 128. 2011.

Epiphytic herbs, to 30 cm tall. Leaf blades oblanceolate to 

obovoid, 6–15 × 1.5–3(–5) cm. Inflorescences 4 to several, fasciculate, 3.5–22 cm long, shorter than or exceeding the leaves, 

many-flowered. Flowers secund, nutant, yellowish-green. Sepals 

subequal, ovate to elliptic-ovate, 2–5 × 1.8–3 mm. Petals trapezoidal. Labellum oblong to rounded.

Found in South America (Venezuela, Colombia, Ecuador, and Peru). In Venezuela, evidently confined to the Andes 

(Mérida, Táchira, and Trujillo; Map 241). Collected along the 

Boconó–Guaramacal road; 2,100–2,300 m.

The status of this record is uncertain since we have not seen 

the voucher (Steyermark & Rabe 97317, VEN) cited by Foldats 

(1970: 158) and subsequently listed by Ortega et al. (1987: 50). 

Foldats (1970: 158) wrote that this species is very close to Stelis oblonga. Jørgensen and León-Yánez (1999: 764), however, 

considered S. striolata to be endemic to Ecuador.

Stelis vulcani Rchb. f., Otia Bot. Hamburg. 1: 19. 1878; Dunsterville and Garay, Venez. Orchids Ill. 4: 324–325. 1966; 

Foldats, Fl. Venez. 15(2): 170–172, fig. 251. 1970; Romero 

and Carnevali, Orchids Venez., 2nd ed., 3: 1056. 2000; 

Fernández, Orquídeas Nat. Táchira 226. 2003; Morillo, 

in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 1: 

306–308, fig. 131. 2011.

Epiphytic or lithophytic herbs, 3–12(–28) cm tall. Leaf 

blades oblong, narrowly oblong to oblong-elliptic, (2.5–)7–12 × 

MAP 240. Stelis pusilla occurrence in Venezuela. MAP 241. Stelis striolata occurrence in Venezuela.

MAP 242. Stelis vulcani occurrence in Venezuela.
NUMBER 100 • 203

(0.6–)2.5–3 cm. Inflorescences 1–4, erect, to 18 cm long, exceeding the leaves, multiflowered. Flowers secund, slightly nutant, 

white, light yellow or green. Sepals unequal, glabrous; dorsal 

sepal elliptic-ovate, 4–6 × 1.7–3.2 mm; lateral sepals 2.8–4 × 

1–1.4 mm. Petals rhomboid. Labellum suborbicular.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, evidently restricted to the Andes (Lara, 

Táchira, and Trujillo; Map 242). In the park, found in cloud forest on both slopes of Guaramacal; 1,850–2,600 m.

Telipogon Kunth

Telipogon Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 269. 1815 [1816]; ibid. 

[qu.] 1: 335. 1815 [1816].

Stellilabium Schltr., Orchideen 530. 1914.

Epiphytic, rarely terrestrial herbs, often growing on twigs or 

thin branches; stems elongated or abbreviated, with monopodial 

or sympodial growth, completely clothed by leaf sheaths; without 

pseudobulbs. Leaves distichous, articulate, elliptic or lanceolate, 

conduplicate, coriaceous or fleshy. Inflorescences terminal or lateral racemes, 1–10-flowered, usually longer than the subtending 

leaves; flowers opening successively (often 1–3 open simultaneously per inflorescence); peduncles often 3- or 4-edged; rachis 

straight or flexuous. Flowers small and inconspicuous or more 

typically medium to large and showy, resupinate or not, with 

widely spreading perianth segments. Floral bracts fleshy, navicular, conspicuous or inconspicuous. Sepals free, subequal, typically 

lanceolate or elliptic. Petals much longer and broader than the 

sepals, free, overlapping and hiding the sepals from view, typically broadly ovate, triangular, or suborbicular, acute or apiculate apically, with 5–15 nerves. Labellum similar to the petals or 

somewhat broader, with 10–30 longitudinal nerves, simple, disk 

naked or provided at the base with a small cushion-like, pubescent or hairy callus; column short and thick, wingless, footless, 

usually bristly or pubescent or both; anther dorsal, erect with reduced partitions; pollinia 4, unequal, with a long, narrow stipe 

and a small hooked viscidium; stigmatic surface apical or subapical, entire; rostellum erect, elongated. Capsules 3-angled to terete 

in cross section. [Epidendroideae: Cymbidieae: Oncidiinae.]

A neotropical genus of ~170 species found in Central America 

(Costa Rica and Panama) and South America (Colombia, Venezuela, Ecuador, Peru, and Bolivia), with a center of diversity in Colombia and Ecuador. Approximately 11 species occur in Venezuela.

References. Dressler (1999); Kränzlin (1919); 

Pridgeon et al. (2009); Sánchez (2002); Senghas (1994b, 1994c); 

Williams et al. (2005).

Telipogon is composed of many narrowly endemic species 

that are often locally common. Plants are usually found at elevations above 2,000 m in constantly cloud-shrouded elfin forest 

and frequently as twig epiphytes.

The number of nerves in the petals (including labellum) is an 

important taxonomic character for distinguishing species.

There is a report, which is not well documented, that the flowers of Telipogon are pseudocopulated by tachinid flies (Tachinidae; 

Maduro in Dressler, 1981: 252). If this can be confirmed, then Telipogon is another example of an orchid being pollinated by deceit.

KEY TO THE SPECIES OF TELIPOGON

1a. Inflorescences many-flowered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. astroglossus

1b. Inflorescences 1–4-flowered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Petals subrotund to narrowly ovoid; ecallose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. andicola

2b. Petals oval to obovate-oblanceolate; callus pilose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. latifolius

Telipogon andicola Rchb. f., Bonplandia (Hannover) 3: 239. 1855; 

Dunsterville and Garay, Venez. Orchids Ill. 5: 304–305. 1972; 

Foldats, Fl. Venez. 15(5): 34–36, fig. 777. 1970 Romero and 

Carnevali, Orchids Venez., 2nd ed., 3: 1067. 2000.

Epiphytic herbs, 8–13 cm tall (including the inflorescence). 

Leaves oblanceolate, 4–6 × 0.7–1.2 cm, bases attenuate, apices 

acute. Inflorescences 2–4-flowered, 1(–2) flowers open at a time. 

Flowers pale green with a maroon-purple center and maroon 

nerves on petals and labellum. Dorsal sepal navicular, 9–14 mm 

long. Petals subrotund to narrowly ovoid, 10–15 × 8–14 mm, 

shortly apiculate, 5–7-nerved.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, known only from the Andes (Táchira 

and Trujillo; Map 243). In the park, found only on the Fila de 

Agua Fría; 2,700–2,800 m.

MAP 243. (Right) Telipogon andicola occurrence in Venezuela.
204 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Telipogon astroglossus Rchb. f., Xenia Orchid. 1: 16, t. 7, fig. 

II, 12–19. 1854.

Cordanthera andina L. O. Williams, Lilloa 6: 244, t. 2. 1941. Stellilabium andinum (L. O. Williams) Garay & Dunst., in Dunsterville 

and Garay, Orchids Venez., ed. 1, 3: 1004. 1979; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 1059. 2000; Dorr et al., Contr. 

U.S. Natl. Herb. 40: 54. 2000 [2001]. Telipogon andinus (L. O. 

Williams) N. H. Williams & Dressler, Lankesteriana 5: 168. 2005, 

non Dodson, 1984.

Stellilabium pogonostalix auct., non (Rchb. f.) Garay & Dunst.; Dunsterville 

and Garay, Venez. Orchids Ill. 2: 336. 1961, pro parte, excluding type 

& fig.; Foldats, Fl. Venez. 15(5): 46–48, fig. 782. 1970, pro parte, excluding type & fig.

Epiphytic herbs. Leaf blades linear to oblong, 0.5–5 × 0.1–1 

cm, apices obtuse to acute. Inflorescences 1(–2), erect or ascending, many-flowered, 5–10 cm long and up to 2× as long as 

the leaves, subterete or only slightly flattened laterally. Flowers 

yellow-green with maroon areas and spots. Sepals ovate, narrowly lanceolate or elliptic, 2.5–4 × 1.2–2 mm. Petals broadly 

oblanceolate, 2.5–4 ×1–2 mm, acute or subulate.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, confined to the Cordillera de Mérida 

(Mérida and Trujillo; Map 244). Collected in cloud forest along 

the Boconó–Biscucuy road; 1,800 m.

We are not certain that this species has been collected 

within the boundaries of the park. Nonetheless, the locality 

(“Boconó–Biscucuy road”) cited by Dunsterville (1986: 336) 

combined with the elevation he reported (1,800 m) suggests 

that this species eventually will be found within the boundaries 

of the park.

The column of Telipogon astroglossus lacks the apically 

branched hairs that characterize its closest relative, T. alticola 

(Dodson & R. Escobar) N. H. Williams & Dressler (=Stellilabium 

pogonostalix sensu Foldats, pro parte). The latter species has entirely yellow or green flowers as opposed to the former species, 

which has flowers with deep maroon areas at the base of the labellum and spots of the same color on petals and sepals. Telipogon 

astroglossus has been confused with T. pogonostalix Rchb. f. 

(≡ S. pogonostalix (Rchb. f.) Garay & Dunst.), which is endemic to 

Ecuador and Peru and has a conspicuously flattened inflorescence.

Telipogon latifolius Kunth, in H. B. K., Nov. Gen. Sp. 1 [fol.]: 

270. 1815 [1816]; ibid. [qu.] 1: 336. 1815 [1816]; Fernández, Orquídeas Nat. Táchira 230. 2003.

Telipogon bruchmuelleri Rchb. f., Linnaea 41: 28. 1877 [1876]; Dunsterville and Garay, Venez. Orchids Ill. 3: 308–309. 1965; Foldats, Fl. 

Venez. 15(5): 37–39, fig. 778. 1970; Ortega et al., BioLlania 5: 50. 

1987; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 1068. 2000 

(“bruechmuelleri”).

Epiphytic herbs; stems short, 2–4 cm long, 3–6-leaved. Leaf 

blades broadly lanceolate, 2.5–6 × 0.4–1.5 cm, bases cuneate, 

apices acute to apiculate. Inflorescences 1- to few-flowered, 4–10 

cm long. Flowers greenish or green-yellow. Dorsal sepal lanceolate, 10–20 mm long. Petals ovate to obovate-oblanceolate, 

12–24 × 6.5–11 mm, bases conspicuously clawed or attenuate, 

apices acuminate, 5–7-nerved.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, this species occurs only in the Andes 

(Mérida, Táchira, and Trujillo; Map 245). Cloud forest along the 

Boconó–Guaramacal road; 1,950 m.

MAP 244. Telipogon astroglossus occurrence in Venezuela. MAP 245. Telipogon latifolius occurrence in Venezuela.
NUMBER 100 • 205

Trichocentrum Poepp. & Endl.

Trichocentrum Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 11. 1838 [1836].

Epiphytic, rarely lithophytic herbs; caespitose or with short 

rhizomes; sympodia terminating in small, subspherical or subcylindric pseudobulbs; pseudobulbs covered by 1 or 2 non-leafbearing scarious sheaths, apically 1-foliate. Leaves conduplicate, 

articulate, elliptic, oblong-elliptic or narrowly obovate, erect, 

arched or subpendulous, fleshy to fleshy-coriaceous. Inflorescences axillary from base of pseudobulbs, 1- to few-flowered racemes, shorter than the leaves, flowers opening in slow gradual 

succession, usually patent, sometimes subpendulous; peduncles 

covered by a few remote, scarious sheaths. Flowers resupinate, 

small to relatively large and showy, relatively short-lived (lasting 2–5 days); perianth segments subfleshy, usually spreading, 

sometimes fragrant, usually white with or without red-purple 

or purplish-brown stripes or spots, sometimes completely pink. 

Sepals similar, free. Petals similar to sepals. Labellum usually 

larger, fleshier, and showier than other perianth segments, adnate 

to the base of the column and parallel to it, usually simple or 

very slightly 3-lobed, basally produced into a conspicuous nectariferous spur of variable length, labellar disk simple or with multiple keels or lamellae; column erect, relatively short and thick, 

winged apically, footless; anther terminal, operculate, united to 

a wide tegula, ± triangular, relatively short; viscidium small; rostellum short; stigma ventral. Capsules ellipsoidal or somewhat 

3-angled, pendent. [Epidendroideae: Cymbidieae: Oncidiinae.]

A subtropical and tropical American genus of 25–50 species 

found in North America (USA), Mexico, Central America, the 

West Indies, and South America (Colombia, Venezuela, Trinidad 

and Tobago, the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, and Argentina). Three species are known from Venezuela.

References. Pridgeon et al. (2009); Pupulin (1995, 

2000); Senghas (1995c); Williams et al. (2001).

Trichocentrum pulchrum Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 

11, t. 115. 1838 [1836]; Dunsterville and Garay, Venez. Orchids Ill. 3: 314–315. 1965; Foldats, Fl. Venez. 15(5): 74–

76, fig. 790. 1970; Romero and Carnevali, Orchids Venez., 

2nd ed., 3: 1075. 2000.

Epiphytic herbs. Leaf blades linear-oblong to oblonglanceolate, 4.5–9 × 1.5–2.5 cm. Inflorescences 1- or 2-flowered. 

Flowers relatively large compared to the size of the plant, white 

with rose tinge. Dorsal sepal 1.8–2.5 cm long. Labellum with a 

yellow and green throat with rose tinge; spur conspicuous, 5–6 

cm long.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru). In Venezuela, known from the Andes (Lara, Mérida, 

Portuguesa, and Trujillo) and the Sierra de Perijá (Zulia; Map 

246). In the park, found on the south slope in sector Paramito–

La Aguadita; 1,800 m.

Amazingly, the first record of the presence of Trichocentrum pulchrum in the park was a sight record by G. Gerlach 

(pers. comm.), who after finding the pollinarium of this species 

on an euglossine bee, Eulaema meriana (Oliver, 1789), that he 

had baited with scent, then looked for and found this epiphytic 

orchid in nearby forest.

Trichopilia Lindl.

Trichopilia Lindl., Edwards’s Bot. Reg. 22: t. 1863. [1 Jun] 1836; Lindl., Intr. 

Nat. Syst. Bot., 2nd ed., 446. [Jul?] 1836.

Pilumna Lindl., Bot. Reg. 30: Misc. p. 73. 1844.

Helcia Lindl., Edwards’s Bot. Reg. 31: Misc. 18. 1845.

Leucohyle Klotzsch, Index Seminum [Berlin] 1854, App.: 1. 1855.

Neoescobaria Garay, Orquideología 7: 194. 1973.

Epiphytic herbs, caespitose, erect, inconspicuous to large 

and showy. Pseudobulbs heteroblastic, unifoliate, usually ovoid 

or fusiform to ovoid, usually strongly compressed laterally, covered by a few scarious sheaths lacking foliar blades. Leaves conduplicate, articulate, usually oblong or elliptic, sessile or with a 

short pseudopetiole, coriaceous. Inflorescences 1 or 2, originating from the bases of the pseudobulbs, racemose, usually shorter 

than the leaves, usually pendulous, rarely suberect or arching, 

1–3(–15)-flowered; peduncles terete to somewhat compressed 

laterally, with a few remote sheaths. Floral bracts conspicuous, shorter than the pedicels. Flowers resupinate, showy, usually opening widely, usually without spurs, yellowish to white 

or creamy green, sometimes with red-brown spots and a yellow 

spot at the base of the labellum. Sepals free or lateral sepals 

shortly fused at the base, undulate-twisted in many species. Petals similar to dorsal sepal. Labellum large and more conspicuous than other segments of the perianth, simple, 3- or 4-lobed, 

lateral lobes enveloping column in a tube, with a conspicuous 

basal claw, the central lobe fused to the center of the column in 

MAP 246. Trichocentrum pulchrum occurrence in Venezuela.
206 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

a group of species (Trichopilia s. str.) or completely free (Helcia

Lindl., Leucohyle Klotzsch, Neoescobaria Garay); disk ecallose 

or consisting of a pair of long keels, aligned below with wings of 

the column near the base of the central lobe, often densely papillose. Column semiterete, straight, ~1⁄2 the length of the labellum; 

anther terminal, operculate, yellow cream, glabrous or papillose, 

subtended by a conspicuous entire or lacerate hood that exceeds 

the anther in length, oblong or subtriangular, caudicles irregular 

shaped; viscidium ventral, oval, apical on the rostellum; pollinia 

2, yellow, waxy, obovoid or pyriform; rostellum shorter than the 

hood of the anther; stigmatic cavity ventral, circular, located just 

below the rostellum. Ovary glabrous, terete. Capsules ellipsoid 

or triangular in cross section. [Epidendroideae: Cymbidieae: 

Oncidiinae.]

A neotropical genus of ~25 species found in Mexico, Central America, the Greater Antilles, and South America (Colombia, Venezuela, Trinidad and Tobago, Guyana, Suriname, 

Ecuador, Peru, Bolivia, and Brazil) but best represented in the 

Andes. Seven species occur in Venezuela.

References. Cavestro (2000); Pridgeon et al. (2009); 

Senghas (1996).

Trichopilia laxa (Lindl.) Rchb. f., Hamburger GartenBlumenzeitung 14: 229. 1858; Dunsterville and Garay, 

Venez. Orchids Ill. 3: 318–319. 1965; Foldats, Fl. Venez. 

15(5): 113–115, fig. 803. 1970; Romero and Carnevali, Orchids Venez., 2nd ed., 3: 1077. 2000. Pilumna laxa Lindl., 

Edwards’s Bot. Reg. 30: Misc. 74. 1844.

Trichopilia sp. A; Dorr et al., Contr. U.S. Natl. Herb. 40: 55. 2000 [2001].

Epiphytic herbs; pseudobulbs 4–5(–12) cm long. Leaf blades 

linear-oblong or lanceolate to elliptic, 14–20(–40) × 2–2.5 cm. 

Inflorescences 5–9-flowered, pendulous, 13–19 cm long. Flowers 

white or pink-tinged, sweetly fragrant. Sepals 3–4 cm long.

Found in the Greater Antilles and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, and Brazil). The West 

Indian record, however, is based on a single poorly preserved 

specimen. In Venezuela, restricted to the Andes (Lara, Mérida, 

Táchira, and Trujillo; Map 247). In the park, found once between El Campamento (below Cerro El Diablo) and Maciegal; 

1,800 m.

Trichosalpinx Luer

Trichosalpinx Luer, Phytologia 54: 393. 1983.

Epiphytic, lithophytic or terrestrial herbs; mostly inconspicuous; usually caespitose but sometimes long-creeping, erect 

to pendulous. Rhizomes abbreviate to long and creeping; stems 

terete, much shorter to several times longer than the leaves, thin 

and wiry to relatively stout, 1-leaved at apex, sometimes apically 

prolific or superposed; enveloped by tubular, ribbed, ± imbricate 

sheaths (lepanthiform sheaths) with oblique, dilated, margined 

ostia, the ribs and margins of the ostia grossly or microscopically 

ciliate to scabrous, rarely glabrous. Leaves conduplicate, articulate, linear-elliptic to orbicular, usually flat, sometimes concave 

or convex, submembranous to coriaceous or fleshy-coriaceous, 

basally sessile to subpetioled, often purple or maroon-tinged. 

Inflorescences originating from apex of stems with an annulus, 

1-flowered to racemose, lax to dense, much shorter to several 

times longer than subtending leaves, erect to pendulous; peduncles remotely sheathed; floral bracts usually inconspicuous; pedicels short and ± stout to filiform and much longer than ovary; 

ovary terete to, more often, obclavate, usually smooth. Flowers usually resupinate, usually inconspicuous, sometimes large 

(relative to plant size), opening widely to campanulate, sometimes cleistogamous; perianth segments hyaline, white, yellowish 

or variously streaked red or purple to completely red or dark 

purple, usually membranous, sometimes produced into tails. Sepals glabrous to pubescent, subsimilar and free, or the laterals 

variously connate into a flat, convex to concave synsepal. Petals 

free, usually smaller than sepals, usually parallel to column. Labellum hinged to column foot or base, simple or 3-lobed, basally 

often with 2 small, retrorse lobes; disk callose or ecallose, usually 

keeled; margins entire to denticulate or fimbriate to ciliate. Column short to long, erect, straight to somewhat arcuate, usually 

with a well-developed foot, sometimes footless, apically often 

winged; anther apical, subapical or ventral, operculate, incumbent, 1-locular; pollinia 2, subspherical to pyriform, unappendaged; clinandrium entire, lobed, fimbriate to ciliate; rostellum, 

ventral to subapical; stigma ventral to subapical. Capsules variable. [Epidendroideae: Epidendreae: Pleurothallidinae.]

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Trinidad and Tobago, the 

Guianas, Ecuador, Peru, Bolivia, and Brazil). Trichosalpinx includes ~150 species and is especially diverse in the Andes; ~20 

species occur in Venezuela.

MAP 247. Trichopilia laxa occurrence in Venezuela.
NUMBER 100 • 207

References. Luer (1983, 1986a, 1997); Pridgeon 

and Chase (2001); Pridgeon et al. (2001b, 2005).

Trichosalpinx species resemble species in some groups of 

Pleurothallis, but the lepanthiform sheaths of the former genus 

permit ready identification.

There is a suggestion that Trichosalpinx may not be monophyletic (Pridgeon et al., 2005), but molecular sampling has not 

included the full range of infrageneric taxa recognized by Luer 

(1997).

KEY TO THE SPECIES OF TRICHOSALPINX

1a. Proliferous herbs (new shoots originating above the base of old stems), prostrate, rooting in the upper internodes of the 

stems; leaves terete or subterete, >15× longer than wide; flowers caudate, appendages >10 mm long . . . . . T. dunstervillei

1b. Caespitose herbs (all new shoots originating from the base of old stems), erect, rooting only at the base of the stems; 

leaves never terete, never >6× longer than wide; flowers caudate or not . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Plants minute; leaf blades <10 mm long; flowers yellow to purple; sepals caudate . . . . . . . . . . . . . . . . . . T. pusilla

2b. Plants slightly larger; leaf blades >10 mm long; flowers variously colored; sepals not caudate . . . . . . . . . . . . . . . . 3

 3a. Plants pendulous; stems with lax and not tightly clasping lepanthiform sheaths; leaf blades elliptic, apices shortacuminate; inflorescences much shorter than the leaves, 5–8-flowered; flowers red-brown to purple, ciliate; sepals 

obtuse; labellum simple, margins ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. dependens

 3b. Plants erect; stems with tightly clasping lepanthiform sheaths; leaf blades elliptic-spathulate to elliptic, apices 

broadly elliptic to obtuse, never acuminate; inflorescences subequal to or exceeding the leaves, 2–5-flowered; 

flowers yellow, glabrous; sepals acute; labellum 3-lobed, margins glabrous . . . . . . . . . . . . . . . . . . T. deceptrix

♦ Trichosalpinx deceptrix Carnevali & I. Ramírez, Harvard Pap. 

Bot. 3: 250, fig. 7. 1998; Romero and Carnevali, Orchids 

Venez., 2nd ed., 3: 1082. 2000; Morillo, in Morillo et al., 

Bot. Ecol. Monocot. Páramos Venez. 1: 316, fig. 1363. 2011.

Epiphytic herbs, 6–8 cm tall. Leaf blades elliptic-spathulate 

to elliptic, 22–30 × 6–8.5 mm; petioles to 4 cm long. Inflorescences 2–5-flowered. Sepals yellow; dorsal sepal deeply concave, 

5 × 2–3 mm. Petals 3 × 0.7 mm, somewhat translucent. Labellum 

3-lobed.

Endemic to Venezuela, where it is found only in the Cordillera de Mérida (Trujillo state; Map 248). Known only from the 

park, where it occurs in cloud forest; ~1,850 m.

Trichosalpinx dependens (Luer) Luer, Phytologia 54: 395. 1983; 

Romero and Carnevali, Orchids Venez., 2nd ed., 3: 1083. 

2000; Fernández, Orquídeas Nat. Táchira 234. 2003. Pleurothallis dependens Luer, Selbyana 3: 94, fig. 150. 1976.

Lepanthes sp. A; Dorr et al., Contr. U.S. Natl. Herb. 40: 51. 2000 [2001].

MAP 248. Trichosalpinx deceptrix occurrence in Venezuela. MAP 249. Trichosalpinx dependens occurrence in Venezuela.
208 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Epiphytic herbs, 3–15 cm tall. Leaf blades elliptic to narrowly elliptic, 30–90 × 10–27 mm; petioles 1–3 mm long. Inflorescences congested, 5–8-flowered. Sepals red-brown to purple; 

dorsal sepal ovate, 2–5 mm long. Petals 1–2 × 0.5–1 mm, translucent. Labellum simple.

Found in South America (Colombia, Venezuela, French Guiana, Ecuador, Peru, and Bolivia). In Venezuela, found only in the 

Andes (Portuguesa and Táchira; Map 249). Cloud forest near La 

Divisoria de la Concepción; 1,700 m.

♦ Trichosalpinx dunstervillei Luer, Monogr. Syst. Bot. Missouri 

Bot. Gard. 64: 55, fig. 43. 1997; Romero and Carnevali, 

Orchids Venez., 2nd ed., 3: 1085. 2000.

Pleurothallis vagans Garay & Dunst., in Dunsterville and Garay, Venez. Orchids Ill. 6: 372–373. 1976, pro parte, excluding type.

Epiphytic herbs, to 15 cm tall. Leaf blades narrowly elliptic, 

20–40 × 2.5–3 mm; petioles 1–2 mm long. Inflorescences fewflowered. Sepals yellow; dorsal sepal concave, 12–14 mm long. 

Petals 2–3 × 0.8 mm, translucent. Labellum 3-lobed.

Endemic to Venezuela, where it is found only in the Cordillera de Mérida (Trujillo state; Map 250). Known from the north 

slope of Guaramacal; 2,350 m.

Trichosalpinx pusilla (Kunth) Luer, Phytologia 54: 397. 1983; 

Morillo, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 1: 321. 2011. Dendrobium pusillum Kunth, in H. 

B. K., Nov. Gen. Sp. [fol.] 1: 286. 1815 [1816]; ibid. [qu.] 

1: 357. 1815 [1816]. Pleurothallis pusilla (Kunth) Lindl., 

Edwards’s Bot. Reg. 28: Misc. 82. 1842; Foldats, Fl. Venez. 

15(2): 371–374, fig. 326. 1970.

Epiphytic, sometimes lithophytic herbs. Leaf blades elliptic to orbicular, 3–8 × 3–5 mm; petioles ~1–2 mm long. Inflorescences 1–3-flowered. Sepals yellow to purple; dorsal sepal 

subcarinate, 4–5.5 mm long. Petals 1.5–2 × 0.7–1 mm, translucent. Labellum 3-lobed.

Found in Central America (Costa Rica) and South America 

(Colombia, Venezuela, Ecuador, Peru, and Brazil). In Venezuela, 

the distribution is disjunct with records from the Andes (Mérida, 

Táchira, and Trujillo) and the Venezuelan Guayana (Amazonas 

and Bolívar; Map 251). Our material is from both slopes of 

Guaramacal; 1,950–2,750 m.

Warreella Schltr.

Warreella Schltr., Orchideen 424. 1914.

Terrestrial herbs, erect; caespitose; pseudobulbs present. 

Roots terete, produced from rhizomes at base of pseudobulbs; rhizomes short, brittle. Pseudobulbs clustered, fusiform or conical, 

homoblastic, with 3–8 internodes, of which 3 or 4 are more developed than the most basal or apical one, when young with 5–8 

distichous leaves and completely hidden by leaf sheaths, at maturity with 3 or 4 leaves concentrated in the upper 1⁄2 of the plant 

and partly naked, covered in part by the fibrous remnants of leaf 

sheaths. Leaves plicate, articulate, membranous, linear-elliptic, 

basally attenuate in a conduplicate pseudopetiole, apically acute. 

Inflorescences erect, raceme emerging from the axils of the lower 

sheaths of the immature pseudobulbs, rachis concentrated in the 

apical 1⁄3, successively flowered, 2–7 flowers open simultaneously; 

peduncles terete with several internodes, each internode subtended 

by lanceolate-elliptic bracts, acute, clasping the peduncle; floral 

bracts narrowly lanceolate or triangular-lanceolate, acute, membranous, shorter than the pedicellate ovaries. Flowers resupinate, 

small, perianth segments widely spreading, white, pink or pale 

violet (often blue at anthesis), nerves often darker purple; labellum white with violet infusion, more intense in the apical 1⁄2; callus 

MAP 250. Trichosalpinx dunstervillei occurrence in Venezuela.

MAP 251. Trichosalpinx pusilla occurrence in Venezuela.
NUMBER 100 • 209

white. Sepals free, somewhat concave, lanceolate to oblanceolate; 

dorsal sepal acute, somewhat reclinate over the column; lateral sepals wider, obtuse to rounded, dorsally carinate, the inner margin 

replicated and folded toward the base. Petals oblanceolate, subacute to rounded. Labellum sessile, articulate with the columnar 

foot, ovate to suborbicular, inconspicuously 3-lobed, apex truncate, somewhat emarginate, basal margins straight and partially 

enveloping the column, apical lobe somewhat reflexed, margins 

undulate; disk with 5 longitudinally flabellate keels originating in 

the labellar base and extending for 1⁄2 the labellar length. Column 

semiterete, recurved, with a conspicuous foot, widened apically; 

anther terminal, operculate, somewhat flattened; pollinia 4 in 2 

unequal pairs, on a triangular stipe and triangular viscidium; stigmatic surface transversely narrowly elliptic. Capsules ellipsoidal. 

[Epidendroideae: Cymbidieae: Zygopetalinae.]

A genus of 3 shade-loving species confined to South America 

(Colombia, Venezuela, and Ecuador); 1 species found in Venezuela.

References. Pridgeon et al. (2009); Senghas (1993a).

Warreella cyanea (Lindl.) Schltr., Orchideen 425. 1914; Dunsterville and Garay, Venez. Orchids Ill. 3: 328–329. 1965; 

Foldats, Fl. Venez. 15(4): 265–267, fig. 666. 1970; Romero 

and Carnevali, Orchids Venez., 2nd ed., 3: 1108. 2000; Llamozas et al., Libro Rojo Fl. Venez. 471. 2003. Warrea cyanea Lindl., Edwards’s Bot. Reg. 30: Misc. 2. 1844.

Terrestrial herbs. Leaf blades 20–50 × 6–10 cm. Flowers 

pale violet. Sepals 1.2–1.5 cm long.

Found in South America (Colombia, Venezuela, and Ecuador). Venezuelan records are from the Andes (Mérida, Táchira, 

and Trujillo), the Sierra de Perijá (Zulia), the Cordillera de la 

Costa (Apure, Aragua, Distrito Federal, and Miranda), and the 

Venezuelan Guayana (Amazonas Map 252). In the park, found 

once in forest below Páramo del Pumar; 1,950–2,150 m.

Poaceae

L. J. DORR, S. MIGUEL NIÑO, AND B. STERGIOS

Annual or perennial, herbaceous or woody plants; caespitose, rhizomatous, stoloniferous, scandent or aquatic. Culms 

(i.e., aerial stems) cylindrical, rarely flattened, hollow, pithy or 

solid; usually branched at the middle and upper nodes; branches 

usually solitary at the nodes (or in some Bambusoideae several 

per node). Leaves alternate, distichous or rarely spirally arranged, 

estipulate; sheaths open or less commonly closed; ligule(s) usually present at junction of sheath and blade; inner ligule present (rarely absent), membranous and/or ciliate; outer ligule 

present in some genera; blades usually linear or lanceolate to 

ovate, flat to inrolled, entire, venation parallel; pseudopetiolate 

or not. Inflorescences compound, terminal and/or axillary, open 

to contracted, composed of few to many discrete spikelets (i.e., 

reduced partial inflorescences) arranged in 1 to several spikes, 

racemes or panicles. Spikelets diverse, but typically composed 

of a rachilla (i.e., axis) bearing at its base 2 persistent or deciduous glumes (i.e., sterile bracts), 0- to several-nerved; rachilla 

above the glumes bearing 1 or more florets. Florets composed of 

a lemma (i.e., subtending bract) and palea (i.e., first appendage 

of the floral axis) that enclose a reduced flower; lemma 1- to 

many-nerved, awned or not; palea usually bicarinate, sometimes 

absent. Flower bisexual or unisexual (rarely gynodioecious), 

perianth reduced, composed of (0) 2 or 3 minute fleshy or scalelike lodicules (i.e., tepals). Stamens (1 or 2)3 or 6 (to many); 

filaments free, rarely connate; anthers basifixed. Ovary superior, 

1-locular, 1-ovulate; placentation subapical to basal. Styles 2. 

Stigmas (1)2 or 3, hispid or plumose. Fruit usually a caryopsis, 

rarely a utricle; seed coat adnate to pericarp; endosperm abundant; embryo small to large.

A cosmopolitan family of ~750 genera and ~10,000 species. 

In Venezuela, there are ~140 genera and ~1,000 species.

References. Campbell (1985); Clark et al. (1995); 

Clayton and Renvoize (1986); Grass Phylogeny Working Group 

(2001); Grass Phylogeny Working Group II (2012); Luces de Febres (1963); Morrone et al. (2012).

KEY TO THE GENERA AND SELECT GROUPS OF POACEAE

1a. Perennial plants; culms usually highly lignified; leaves dimorphic, culm and foliage leaves distinct . . . . . . . . . . . . . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bambusoideae (see separate key)

1b. Annual or perennial plants; culms herbaceous or subwoody (i.e., easily crushed with the fingers); leaves not dimorphic 

(except Dichanthelium where basal and culm leaves are distinct) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

MAP 252. Warreella cyanea occurrence in Venezuela.
210 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

2a. Spikelets disarticulating above the glumes; empty glumes remaining attached to the pedicels . . . . . . . . . . . . . . . . 3

 3a. Spikelets with 1 floret, additional rudimentary florets lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. One or both glumes shorter than the lemma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sporobolus

 4b. Both glumes longer than the lemma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 5a. Plants delicate, <40 cm tall; panicles delicate, open, and plumose; paleas absent or minute . . . Agrostis

 5b. Plants robust, >40 cm tall; panicles dense to closed; paleas well developed, ± equal in length to lemmas 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calamagrostis

 3b. Spikelets with 2 to many florets, all well developed or several rudimentary (reduced or sterile) . . . . . . . . . . . 6

 6a. Plants gynodioecious; culms to 1.5 m tall; leaf sheaths often curling and disintegrating with age; inflorescences plumose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cortaderia

 6b. Plants not gynodioecious; culms usually <1 m tall; leaf sheaths not as above; inflorescences not plumose . . . . 7

 7a. Glumes subequal, equal to or longer than the spikelet (not including awns) . . . . . . . . . . . . Danthonia

 7b. Glumes unequal, 1 or both much shorter than the spikelet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

 8a. Lemmas with 3 conspicuous nerves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eragrostis

 8b. Lemmas with 5 or more inconspicuous nerves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

 9a. Leaf sheath margins joined for at least 1⁄3 of their length . . . . . . . . . . . . . . . . . . . . . . . Bromus

 9b. Leaf sheath margins usually free and overlapping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

 10a. Lemmas awnless, acute or blunt; leaf blades with navicular apices . . . . . . . . . . . . . . Poa

 10b. Lemmas awned or conspicuously acuminate; leaf blades with acuminate apices . . . Festuca

2b. Spikelets disarticulating individually below the glumes or falling in aggregates; empty glumes not remaining attached 

to pedicels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

11a. Spikelets disarticulating individually without other structures attached . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12a. Spikelets dorsally compressed or terete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

 13a. Spikelets subtended or concealed by long and conspicuous silky hairs or bristles . . . . . . . . . . . . . . . . . . . 14

 14a. Spikelets subtended by 1 to several bristles; bristles antrorsely scabrous . . . . . . . . . . . . . . . . . . . Setaria

 14b. Spikelets concealed by long, silky hairs; hairs not antrorsely scabrous . . . . . . . . . . . . . . . . Eriochrysis

 13b. Spikelets glabrous or pubescent with short and inconspicuous hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

 15a. Inflorescence 1-pinnate, lateral racemes not branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

 16a. Backs of fertile lemmas and upper glumes turned toward rachis . . . . . . . . . . . . . . . . . . Paspalum

 16b. Backs of fertile lemmas and upper glumes turned away from rachis . . . . . . . . . . . . . . . Axonopus

 15b. Inflorescence 2- or 3-pinnate, lateral racemes (at least the lower ones) branched . . . . . . . . . . . . . . . 17

 17a. Spikelets globose, subglobose or obovoid, or, if ellipsoid, then inserted obliquely on the pedicel . . . . 18

 18a. Ligules a row of hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Isachne

 18b. Ligules a membrane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lasiacis

 17b. Spikelets ellipsoid or lanceoloid, not inserted obliquely on the pedicel . . . . . . . . . . . . . . . . . . . 19

 19a. Upper lemma with 2 winglike appendages adnate to the base of the lemma and free above, 

sometimes appendages reduced to swellings . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ichnanthus

 19b. Upper lemma without winglike appendages or swellings . . . . . . . . . . . . . . . . . . . . . . . . . 20

 20a. Glumes subequal, ± equal in length to spikelet . . . . . . . . . . . . . . . . . . . . . . . Homolepis

 20b. Glumes unequal, the lower glume conspicuously shorter than the spikelet . . . . . . . . 21

 21a. Terminal inflorescences with chasmogamous spikelets; axillary inflorescences with 

cleistogamous spikelets; basal and culm leaves different in size and shape . . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dichanthelium

 21b. Terminal and axillary inflorescences with chasmogamous spikelets only; basal and 

culm leaves similar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

 22a. Lower lemmas with 1 or 2(3) pairs of crateriform or ocellate glands . . . . . .

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ocellochloa

 22b. Lower lemmas without glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rugoloa

12b. Spikelets compressed laterally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

 23a. Leaves densely viscid-hairy (our species); lower lemma awned; awn 5–8 mm long . . . . . . . . . . . . . . Melinis

 23b. Leaves not viscid; lower lemma awnless or awn <5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

 24a. Glumes longer than the floret . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

 25a. Glumes scabrous, awnless or minutely awn-tipped; awn <0.4 mm long . . . . . . . . . . . . . . . Cinna

 25b. Glumes hispidulous, awned; awns 1–2(–4) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . Polypogon

 24b. Glumes (at least the lower one) shorter than the floret . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ichnanthus
NUMBER 100 • 211

11b. Spikelets (all or some) falling in clusters of 2 or more, or falling while attached to rachis internodes . . . . . . . . . . . . 26

26a. Spikelets surrounded by a fascicle of bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cenchrus

26b. Spikelets not surrounded by a fascicle of bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

 27a. Inflorescences complex plumose or cylindrical panicles with numerous rames (i.e., sessile-pedicellate spikelet 

pairs and internodes forming dispersal units) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

 28a. Spikelets visible between a pubescence of white or whitish-cream to cream hairs . . . . . . Andropogon

 28b. Spikelets completely obscured by a pubescence of chestnut or coppery-brown hairs . . . . . Eriochrysis

 27b. Inflorescences flabellate (our species) with 12–20 rames . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arthraxon

KEY TO THE GENERA OF BAMBUSOIDEAE

1a. Aerial culms never branched; leaf blades >50 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chusquea

1b. Aerial culms branched; leaf blades <50 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Branch complement at midculm nodes consisting of 3 to many branchlets; leaf blades not reflexed . . . . . . . . . . . 3

 3a. Internodes of culms solid, sometimes becoming fistulose (i.e., reedlike or hollow) . . . . . . . . . . . . . . Chusquea

 3b. Internodes of culms hollow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Primary element of the branch complement at midculm nodes terete, segmented, and diverging from the 

culm; second-order branches fasciculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arthrostylidium

 4b. Primary element of the branch complement at midculm nodes flat, unsegmented, appressed and adnate to 

the culm surface; second-order branches flabellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Didymogonyx

2b. Branch complement at midculm nodes usually consisting of only 1 branchlet; leaf blades reflexed . . . . . Aulonemia

Agrostis L.

Agrostis L., Sp. Pl. 61. 1753, nom. cons.

Perennial or less commonly annual plants; caespitose, sometimes rhizomatous or stoloniferous. Culms erect, ascending, decumbent or prostrate; unbranched or rarely sparsely branched. 

Leaves cauline and basal, few to many; sheaths open; ligules 

membranous, usually erose to lacerate; blades linear, flat, folded 

or involute, rigid or not. Inflorescences terminal panicles, open or 

contracted; primary panicle branches whorled or not. Spikelets 

1-flowered, laterally compressed, with or without a barren rachilla 

extension; flowers perfect. Disarticulation above the glumes, below 

the florets. Glumes equal or subequal, rarely unequal, membranous, 1(–3)-nerved, acute, acuminate to awned. Lemmas usually 

shorter than the glumes, hyaline, faintly 3–5-nerved, apex blunt 

or narrow, sometimes toothed or awned. Paleas absent or minute. 

Lodicules 2. Anthers (1) 3. Ovary glabrous. Styles 2. Caryopses 

subterete, ellipsoid; pericarp adherent. [Pooideae.]

A cosmopolitan genus of ~220 species found in temperate and 

cool regions. In the Americas, found in North America (Greenland, 

Canada, and USA), Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Chile, Paraguay, Argentina, and Uruguay). In Venezuela, ~15 

species are known from the Andes and the Cordillera de la Costa.

References. Rúgolo de Agrasar and Molina (1992, 

1993, 1997); Tucker (1996).

Agrostis is closely related to Calamagrostis Adans. and Deyeuxia Clarion ex P. Beauv. The relationship of these three genera, 

part of an intergrading complex of incompletely separated entities, is discussed under Calamagrostis.

KEY TO THE SPECIES OF AGROSTIS

1a. Awns on lemmas >2 mm long; leaves mostly basal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Panicles linear, narrow, green; lemmas 1.9–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pittieri

2b. Panicles broadly ovate to lanceolate, spreading, purplish; lemmas 2–2.5 mm long . . . . . . . . . . . . . . . . A. mertensii

1b. Awns on lemmas <2 mm long or absent; leaves mainly cauline or basal and cauline . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Panicles broadly ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Panicle branches divided below the middle; rachilla and pedicels glabrescent . . . . . . . . . . . . . . . . A. perennans

 4b. Panicle branches divided above the middle; rachilla and pedicels notably scabrous . . . . . . . . . . . . . . A. scabra

3b. Panicles linear or ovate to narrowly ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 5a. Glumes 3-nerved, 3.5–3.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. meridensis

 5b. Glumes 1-nerved, 3–3.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. venezuelana

Agrostis meridensis Luces, Bol. Soc. Venez. Ci. Nat. 15: 11, fig. 

7. 1953.

Perennial plants; caespitose. Culms 45–70 cm tall, ascending; nodes glabrous. Leaves mostly cauline; sheaths pubescent; 

ligules 1–2 mm long; blades linear, 7–15 cm × 3–7 mm, scabrous 

above and below. Inflorescences ovate to narrowly ovate, 10–15 × 

4–9 cm long, open, green or purplish; branches whorled at nodes. 

Spikelets elliptic, 3.5–3.8 mm long. Glumes subequal, lanceolate, 
212 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

3.5–3.8 mm long, 3-nerved; lower glume hispidulous. Lemmas 

ovate, 2.2–2.5 mm long, 5-nerved, awnless. Palea minute.

Endemic to Venezuela, where it is known from the Cordillera de Mérida (Mérida and Trujillo) and the Cordillera de la 

Costa (Anzoátegui; Map 253). Páramo and subpáramo near the 

Laguna del Pumar; 3,000 m.

Agrostis mertensii Trin., Linnaea 10: 302. 1836.

Perennial plants; caespitose or rhizomatous. Culms 10–60 

cm tall, erect or ascending. Leaves mostly basal or basal and 

cauline; sheaths glabrous; ligules 1–2(–6) mm long; blades flat, 

occasionally involute, (2.5–)5–10 cm × 1–3 mm, smooth or 

scaberulous. Inflorescences broadly ovate to lanceolate, 5–15 × 

1.5–5(–16) cm, spreading, purplish; branches (1–)2–5 per lower 

nodes. Spikelets lanceolate to narrowly ovate, 2.5–3 mm long. 

Glumes unequal, elliptic to lanceolate, 2.5–3 mm long, 1-nerved, 

lower glume scaberulous. Lemmas elliptic, 2–2.5 mm long, 

5-nerved, awned; awns 3–4.4 mm long. Palea absent or minute.

Found in North America (Greenland, Canada, and USA), 

Mexico, and South America (Venezuela, Ecuador, Peru, Bolivia, 

Brazil, Chile, and Argentina). In Venezuela, known only from 

the Cordillera de Mérida (Trujillo; Map 254). In the park, found 

in open areas of páramo and subpáramo near the Páramo de 

Guaramacal and the Laguna del Pumar; 3,000–3,100 m.

Agrostis perennans (Walter) Tuck., Amer. J. Sci. Arts 45: 44. 

1843; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 553, fig. 9. 2010. Cornucopiae perennans

Walter, Fl. Carol. 74. 1788 (“Cornucopiæ”).

Agrostis alba auct., non L.; Dorr et al., Contr. U.S. Natl. Herb. 40: 56. 2000 

[2001] (as to Cuello et al. 1267).

Agrostis subrepens auct., non (Hitchc.) Hitchc.; Dorr et al., Contr. U.S. Natl. 

Herb. 40: 56. 2000 [2001].

Perennial plants; caespitose. Culms (10–)50–120 cm tall, 

erect or decumbent; sometimes rooting at the nodes. Leaves 

mostly cauline; sheaths glabrous; ligules 2–5 mm long; blades 

flat, rarely involute, 5–13(–20) cm × 2–5 mm, scaberulous. Inflorescences broadly ovate, 12–23(–30) × 2.5–11 cm, open and 

delicate, pale green or more commonly purple; branches 3–11 

per node. Spikelets lanceolate to narrowly ovate, 2.5–3 mm long. 

Glumes unequal, ovate, 2–3 mm long, 1-nerved, nerves scabrous. 

Lemmas ovate, 1.7–2(–2.5) mm long, 5-nerved, awnless (rarely 

awned). Palea minute or absent.

MAP 254. Agrostis mertensii occurrence in Venezuela.

MAP 253. Agrostis meridensis occurrence in Venezuela.

MAP 255. Agrostis perennans occurrence in Venezuela.
NUMBER 100 • 213

Found in North America (Canada and USA), Mexico, Central America, the Greater Antilles, and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Chile, and Argentina). In 

Venezuela, found in the Andes (Barinas, Lara, Mérida, Táchira, 

and Trujillo), the Cordillera de la Costa (Aragua, Distrito Federal, Miranda, and Sucre), and possibly the Sierra de Perijá 

(Zulia; Map 255). In the park, found in open areas of cloud 

forest, subpáramo, and páramo on both slopes of Guaramacal; 

2,000–3,100 m.

Agrostis pittieri Hack., Oesterr. Bot. Z. 52: 60. 1902.

Perennial plants; caespitose, densely clumped. Culms 

30–70 cm tall, erect. Leaves mostly basal; sheaths glabrous or 

scaberulous apically; ligules 2.5–3.5 mm long; blades filiform, 

involute, 7–15 cm × 1–1.5 mm, coarsely ridged above, mostly 

scaberulous below. Inflorescences linear, 9–11 × 1–3 cm, narrow but loose, slender, erect, green; branches 2 to many per 

node. Spikelets elliptic, 3–4 mm long. Glumes subequal, oblong, 2.8–3.9 mm long, 1-nerved, keels scabrous. Lemmas 

ovate, 1.9–2 mm long, faintly nerved, awned; awns 2–3 mm 

long. Palea minute.

Found in Central America (Costa Rica) and South America 

(Venezuela). In Venezuela, known only from higher elevations in 

the Cordillera de Mérida (Mérida and Trujillo; Map 256). In the 

park, found in páramo between the Páramo de Guaramacal and 

the Laguna del Pumar; 2,900–3,000 m.

Our material is referred to this species with some doubt. Although Agrostis pittieri is accepted by Zuluoaga et al. in Hokche 

et al. (2008) as occurring in Venezuela, Pohl (1980) and Morales 

in Hammel et al. (2003b) consider the species to be endemic to 

Costa Rica.

• Agrostis scabra Willd., Sp. Pl. 1(1): 370. 1797; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 554, fig. 

10. 2010.

Agrostis alba auct., non L.; Dorr et al., Contr. U.S. Natl. Herb. 40: 56. 2000 

[2001] (as to Stergios et al. 16114).

Agrostis stolonifera auct., non L.; Bono, Cat. Fl. Vert. Occid. Cordillera 

Mérida 129. 2010.

Annual or perennial plants; caespitose. Culms 30–60 cm 

tall, erect. Leaves mostly cauline; sheaths usually glabrous, sometimes scabridulous; ligules 1.5–4 mm long; blades flat or involute, 4–14(–20) cm × 1–3 mm, scaberulous, especially margins. 

Inflorescences broadly ovate, 7–30 × 5–20 cm, open and spreading, greenish or reddish; branches 2–7 per node (entire panicle 

often detaching at base at maturity). Spikelets lanceolate, 2–3.5 

mm long. Glumes unequal, lanceolate, 2–3.5 mm long. Lemmas oblong, 1–1.2(–2.5) mm long, 5-nerved, muticous, awned 

or not; awns minute, 0.2–0.3 mm long. Palea absent or minute.

Found in North America (Greenland, Canada, and USA), 

Mexico, and the northern Pacific Rim in Asia; introduced in the 

Greater Antilles, South America (Venezuela, Chile, and Argentina), and Europe. In Venezuela, only known from the Cordillera 

de Mérida (Mérida and Trujillo; Map 257). In the park, generally 

in disturbed sites and often on road or trail cuts; 2,000–2,600 m.

Agrostis venezuelana Mez, Repert. Spec. Nov. Regni Veg. 18: 4. 

1922; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 559, fig. 14. 2010.

Perennial plants. Culms 30–40 cm tall. Leaves mostly basal; 

sheaths glabrous; ligules 1–2 mm long; blades linear, ~15 cm × 

2–3 mm, ± glabrous. Inflorescences linear to narrowly ovate, 

8–10 × ~2 cm; branches whorled at nodes. Spikelets elliptic, 

3.5–3.8 mm long. Glumes equal, lanceolate, 3–3.3 mm long, 

MAP 256. Agrostis pittieri occurrence in Venezuela.

MAP 257. Agrostis scabra occurrence in Venezuela.
214 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

1-nerved; lower glume scaberulous. Lemmas elliptic, ~1.8 mm 

long, 5-nerved. Palea absent or minute.

Endemic to Venezuela, where it is found in the Andes 

(Mérida, Táchira, and Trujillo) and the Cordillera de la Costa 

(Distrito Federal and Miranda; Map 258). In the park, found 

in open areas of páramo and subpáramo near the Páramo de 

Guaramacal and the Laguna del Pumar; 2,800–3,000 m.

Andropogon L.

Andropogon L., Sp. Pl. 1045. 1753, nom. cons.

Perennial or less commonly annual plants; caespitose, rarely 

rhizomatous. Culms erect or ascending, much-branched above 

the middle. Sheaths slightly inflated; ligules membranous, eciliate 

or ciliolate; blades linear. Inflorescences 1 or more terminal and 

axillary racemes, often forming complex, compound panicles; racemes composed of numerous rames; peduncles concealed by leaf 

sheaths; disarticulation in the rames below the sessile spikelets. 

Pedicellate spikelets usually vestigial or absent. Sessile spikelets 

bisexual (pistillate), awned or not (our species). Glumes dissimilar; lower glume 2-keeled, flattened or concave, unnerved between 

the keels or sometimes 2–9-nerved; upper glume strongly keeled, 

1–3-nerved, usually awnless. Lodicules 2. Anthers 1, 3(2). Caryopses linear or narrowly ovoid; pericarp adherent. [Panicoideae.]

A cosmopolitan genus of 100–120 species found in tropical 

and temperate regions. In the Americas, found in North America 

(Canada and USA), Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Trinidad and Tobago, the 

Guianas, Ecuador, Peru, Bolivia, Brazil, Chile, Paraguay, Argentina, and Uruguay). In Venezuela, 16 species have been reported.

References. Campbell (1983); Mathews et al. 

(2002); Skendzic et al. (2007).

The morphological limits of Andropogon, a large and variable genus, are problematic. Mathews et al. (2002), in a molecular analysis of the Andropogoninae, suggested that Andropogon

could prove to be paraphyletic. Skendzic et al. (2007) arrived at 

a similar conclusion, i.e., that Andropogon is not monophyletic, 

but these hypotheses largely remain untested, and generic circumscriptions and relationships within the tribe will not be resolved 

without extensive molecular sampling on a worldwide scale.

Andropogon bicornis L., Sp. Pl. 1046. 1753 (“bicorne”), nom. 

cons.; Davidse, in Berry et al., Fl. Venez. Guayana 8: 19. 

2004.

FIGURE 26B

Andropogon bicornis var. burchellii Hack., in Martius, Fl. Bras. 2(3): 285. 

1883; Davidse, in Berry et al., Fl. Venez. Guayana 8: 19. 2004.

Perennial plants; caespitose, in large clumps. Culms 1–1.5

(–2.5) m tall, branching above the middle. Sheaths densely overlapping below, shorter than internodes above, glabrous; ligules 

~1 mm long; blades linear, 30–50 cm × 3–6 mm, margins scabrous, glabrous above and below. Inflorescence a large, plumose, whitish, obovate mass composed of numerous repeatedly 

branching axillary branches that terminate in peduncles bearing 

1–3 bladeless sheaths and a pair of rames. Rames divergent, 2.5–

3.5 cm long. Spikelets paired (pedicellate and sessile). Pedicellate 

spikelets rudimentary or absent; peduncles 3.5–4 mm long. Sessile (fertile) spikelets ovate, 3–3.5 mm long, acute; lower glume 

flattened, nerveless; upper glume naviculiform, 2–2.5 mm long, 

1-nerved; lower (sterile) lemma resembling lower glume, hyaline, 

nerveless; upper (fertile) lemma smaller, 1.5–2 mm long, hyaline, 

nerveless. Anthers 3. Caryopses 1.5–2 mm long.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Trinidad and Tobago, the 

MAP 258. Agrostis venezuelana occurrence in Venezuela.

MAP 259. Andropogon bicornis occurrence in Venezuela.
NUMBER 100 • 215

Guianas, Ecuador, Peru, Bolivia, Brazil, Chile, Paraguay, and 

Argentina); also collected in Florida (USA) but probably not 

established in North America. Widespread in Venezuela (Amazonas, Anzoátegui, Apure, Aragua, Barinas, Bolívar, Cojedes, 

Delta Amacuro, Distrito Federal, Falcón, Guárico, Lara, Mérida, 

Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, 

Trujillo, Yaracuy, and Zulia; Map 259). In disturbed, open areas 

of the park, especially along the track from El Campamento to 

Qda. Honda; 1,900–2,000 m.

• Arthraxon P. Beauv.

Arthraxon P. Beauv., Ess. Agrostogr. 111. 1812.

Annual or perennial herbs; caespitose. Culms ascending 

or decumbent, often rooting at nodes. Sheaths open, slightly 

inflated, glabrous to pubescent; ligules membranous, fimbriate 

or ciliate; blades flat, ovate, ovate-oblong or ovate-lanceolate, 

apex acute to acuminate, base cordate, ± amplexicaul, glabrous 

or pubescent. Inflorescences terminal and axillary panicles, 

panicles subdigitate, often flabellate, clusters of rames; disarticulation in the rames, beneath the sessile spikelets. Spikelets in 

sessile-pedicellate pairs or appearing solitary and sessile, pedicels 

greatly reduced and lacking spikelets. Sessile spikelets bisexual, 

florets 2; glumes equal or subequal; lower florets sterile, reduced 

to an unarmed lemma; upper florets bisexual, awned; anthers 2 

or 3. Pedicellate spikelets absent or rudimentary. Caryopses ellipsoid or fusiform. [Panicoideae.]

Seven species found in tropical and subtropical Africa, Asia, 

and Australia, 1 of which is introduced and naturalized in the 

Americas.

References. van Welzen (1981, 1993).

• Arthraxon hispidus (Thunb.) Makino, Bot. Mag. (Tokyo) 26: 

214. 1912. Phalaris hispida Thunb., in Murray, Syst. Veg., 

ed. 14, 104. 1784.

Alectoridia quartiniana A. Rich., Tent. Fl. Abyss. 2: 448, t. 99 [as “102” 

in text]. 1851. Arthraxon quartinianus (A. Rich.) Nash, N. Amer. Fl. 

17(2): 99. 1912.

The nominate variety is the only one known from Guaramacal and Venezuela.

• Arthraxon hispidus var. hispidus

Annual herbs; caespitose. Culms 50–75 cm long, often 

decumbent, rooting at nodes; nodes pubescent. Sheaths much 

shorter than internodes, glabrous with ciliate margins; ligules 

ciliate; blades ovate to ovate-lanceolate, 1–6 cm × 2–6 mm, glabrous above and below or hispidulous below, margins ciliate. 

Inflorescences terminal, flabellate, 1.5–5 cm long, purplish, with 

12–20 rames; rame internodes pilose with long erect hairs. Sessile spikelets: glumes 3–6 mm long; lower lemmas minute; upper 

lemmas awned, awns 0–12 mm long, usually twisted below, exserted 3–5 mm; paleas absent. Pedicellate spikelets absent.

Native to tropical Africa, Asia, and Australia, but now naturalized throughout the world except for Europe and Antarctica. In the Americas, found in North America (USA), Central 

America, the West Indies, and South America (Venezuela and 

Ecuador). In Venezuela, known from scattered localities in the 

Andes (Lara, Portuguesa, and Trujillo) and the Cordillera de la 

Costa (Guárico; Map 260). In the park, known from the Laguna 

de Aguas Negras; 1,900 m.

The introduction and spread of Arthraxon hispidus var. hispidus in the Americas was traced by Dorr and Niño (2001); it 

first appeared in North America in the 1870s, the West Indies in 

the 1890s, Central America in the 1930s, and South America in 

the 1970s. Venezuelan collections (Dorr and Niño, 2001; Meier, 

2005) were the first to document its occurrence in South America, where it has the potential to become an invasive weed.

Arthrostylidium Rupr.

Arthrostylidium Rupr., Bambuseae 27. 1839 [preprint]; Mém. Acad. Imp. 

Sci. Saint-Pétersbourg, sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 5(2, 

Bot.): 117. 1840.

Small to medium-sized woody bamboos; caespitose; rhizomes pachyform. Culms unarmed, erect at base, scandent or 

climbing above; primary element of the branch complement at 

midculm nodes terete, segmented, and diverging from the culm; 

second-order branches fasciculate; internodes subequal, hollow. 

Culm leaves: well developed or not; sheaths without auricles, 

apices fimbriate or not, fimbriae inconspicuous to conspicuous; 

blades erect, rarely reflexed, bases continuous with the apex 

of sheaths (not constricted), persistent or deciduous. Foliage 

leaves: sheaths with fimbriate apices, fimbriae inconspicuous to 

conspicuous; ligules internal and external, membranous; oral 

setae inconspicuous to prominent, rarely absent; blades linear to lanceolate or ovate to elliptic, midnerve not prominent; 

MAP 260. Arthraxon hispidus var. hispidus occurrence in 

Venezuela.
216 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

pseudopetiolate. Inflorescences spicate or racemose, terminal; 

rachises straight, flexuous or strongly zigzag. Spikelets subsessile, bisexual, 2- to many-flowered. Glumes 1 or 2, shorter than 

spikelets; lowermost florets usually sterile; functional florets 1 to 

several; florets disarticulating and falling attached to the rachilla 

internode; uppermost 1 to several florets successively reduced 

and sterile; lemmas and paleas subequal; paleas with unwinged 

keels; lodicules (0) 3; anthers 3; stigmas 2. [Bambusoideae.]

As presently construed, a neotropical genus of ~32 species 

found in Mexico, Central America, the West Indies, and South 

America (Colombia, Venezuela, Trinidad and Tobago, Guyana, 

Suriname, Ecuador, Peru, Bolivia, and Brazil). Seven species 

occur in Venezuela.

References. Clark and Ely (2011); Judziewicz and 

Clark (1993); Judziewicz et al. (1999); Tyrrell et al. (2012).

This genus has been a catchall among arthrostylidioid bamboos, and as a consequence, the current concept is polyphyletic 

(Judziewicz and Clark, 1993; Tyrrell et al., 2012). Delineation of 

a monophyletic genus likely will involve transferring most species 

of Rhipidocladum McClure (excepting sect. Didymogonyx L. G. 

Clark & Londoño) to Arthrostylidium and the exclusion of a number of morphologically anomalous species of Arthrostylidium.

KEY TO THE SPECIES OF ARTHROSTYLIDIUM

1a. Rachises of mature inflorescences straight; spikelets 2–3 cm long; culms 10–25 mm in diameter; internodes 35–50 cm 

long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pubescens

1b. Rachises of mature inflorescences flexuous or zigzag; spikelets 1–1.7(–3) cm long; culms 3–5(–12) mm in diameter; internodes 18–26 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. venezuelae

Arthrostylidium pubescens Rupr., Bambuseae 29, t. 4, fig. 14. 

1839 [preprint]; Mém. Acad. Imp. Sci. Saint-Pétersbourg, 

sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 5(2, Bot.): 119, t. 4, 

fig. 14. 1840; Judziewicz, in Berry et al., Fl. Venez. Guayana 

8: 36. 2004.

Woody bamboos. Culms scandent above, 5–8 m long, 

5–25 mm in diameter, almost always wider than the secondary 

branches; internodes 35–50 cm long, glabrous or slightly scabrous; nodes with 3–20 or more secondary branches; branches 

subequal or with 1 more robust than the others. Culm leaves: 

20–47 cm long, scabrous, deciduous; sheaths 8–17 cm long; 

blades narrowly triangular, 11–36 cm long, erect, persistent, 

apices acuminate, bases with tiny fimbriate auricles on 1 side 

only. Foliage leaves: 2–12 per branch; sheaths glabrous; ligules 

short, oral setae 3–10 mm long; blades ovate to lanceolate, 

10–15 × 1–2 cm, glabrous above but usually 2 main nerves along 

1 margin scabrous, glabrous below except short-pilose at base; 

pseudopetioles 2–4 mm long. Synflorescences spicate, terminal, 

10–15 cm long, with 6–10 spikelets; rachises straight, glabrous. 

Spikelets appressed to rachis, 2–3 cm long, with 2–4 florets; 

lower glumes 4–7 mm long, 3-nerved; upper glumes 7–10 mm 

long, 3(–5)-nerved; lemmas often ciliolate apically, 9–11-nerved; 

paleas usually equal to or slightly longer than lemmas.

Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, and Trinidad and Tobago). Widespread in Venezuela (Aragua, Bolívar, Distrito Federal, Falcón, Mérida, Miranda, Táchira, Trujillo, and Yaracuy; 

Map 261). In the park, found in forest understory on the north 

slope; 1,950–2,300 m.

This species is distinguished easily from the following one 

when fertile but is confused with it when sterile.

Arthrostylidium venezuelae (Steud.) McClure, J. Wash. Acad. 

Sci. 32: 172. 1942; Judziewicz, in Berry et al., Fl. Venez. 

Guayana 8: 36. 2004. Chusquea venezuelae Steud., Syn. Pl. 

Glumac. 1: 337. 1854.

Woody bamboos. Culms scandent or climbing above, 3–10 

m long, 3–5(–12) mm in diameter, almost always the same thickness as the branches or slightly thicker; internodes 18–26 cm long, 

glabrous; nodes with 10–18(–20) secondary branches; branches 

subequal or with 1 more robust than the others. Culm leaves 

(poorly known): ~24 cm long, glabrous, deciduous?; sheaths 9 cm 

long; blades 15 cm long, erect, persistent. Foliage leaves: 2–6 per 

branch; sheaths with apical appendices 3–5 mm long, ± glabrous; 

ligules short, oral setae 3–6 mm long, rarely absent; blades ovate 

to lanceolate, 4–15 cm × 6–18 mm, glabrous toward the apex, 

with scattered pubescence at base and on margins; pseudopetioles 

MAP 261. Arthrostylidium pubescens occurrence in Venezuela.
NUMBER 100 • 217

1–4 mm long. Synflorescences spicate or racemose, terminal, 

5–10 cm long, with 5–8 spikelets; rachises flexuous or strongly 

zigzag at maturity, glabrous. Spikelets 1–1.7(–3) cm long, with 

4–8 florets; lower glumes 3–4 mm long, 3–5-nerved; upper 

glumes 5.5–6.5 mm long, 5–7-nerved; lemmas glabrous to pilose, 

9–11-nerved; paleas ± equal to lemmas or slightly shorter.

Found in Mexico, Central America, the Lesser Antilles, and 

South America (Colombia, Venezuela, Guyana, Peru, Bolivia, 

and Brazil). In Venezuela, very common in forest and disturbed 

areas in the Andes (Lara, Mérida, Portuguesa, and Trujillo), 

the Cordillera de la Costa (Aragua, Distrito Federal, Falcón, 

Miranda, Sucre, and Yaracuy), and the Venezuelan Guayana 

(Amazonas and Bolívar; Map 262). In the park, found in forest 

understory on both slopes; 1,600–2,300 m.

Aulonemia Goudot

Aulonemia Goudot, Ann. Sci. Nat., Bot., sér. 3, 5: 75. 1846.

Small to medium-sized bamboos; caespitose; rhizomes short, 

pachyform. Culms unarmed, erect or scandent, hollow; midculm 

nodes usually bearing a single, dominant branch; internodes subequal; secondary branches solitary or numerous, the same thickness as the main culm or slightly thinner. Culm leaves: sheaths 

persistent, apices fimbriate, not auriculate, margins fimbriate or 

not; ligules internal and external, oral setae present or absent; 

blades almost always deciduous. Foliage leaves: sheaths persistent, apices fimbriate, margins fimbriate or not; ligules internal 

and external, oral setae present or absent; blades pseudopetiolate, linear, lanceolate or ovate. Inflorescences paniculate, open, 

terminal. Spikelets pedicellate with 2–23 florets; uppermost floret reduced and sterile; disarticulation above the glumes. Lower 

glumes small, 1–3-nerved; upper glumes obtuse, 5–7-nerved; 

lowermost floret sometimes sterile; lemmas 7–9-nerved, mucronate or awned; paleas 2-keeled, not winged; lodicules 3; anthers 

3; stigmas 2. [Bambusoideae.]

A neotropical genus of ~60 species found in Mexico, Central America, and South America (Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Bolivia, and Brazil). Nine species 

occur in Venezuela, all in mountainous regions above 1,800 m 

elevation.

References. Calderón and Soderstrom (1980); 

Clark and Ely (2011); Judziewicz et al. (1999); McClure (1973); 

Tyrrell et al. (2012).

Aulonemia can be recognized by its reflexed leaf blades, 

well-developed apical and sometimes marginal fimbriae on leaf 

sheaths, paniculate inflorescences, and spikelets with two to 

many fertile florets. Molecular data (Tyrrell et al., 2012) suggest that Aulonemia as presently circumscribed is polyphyletic; 

it clearly shows an affinity to Colanthelia McClure & E. W. Sm., 

and at least one species described as Aulonemia is more closely 

related to the monotypic Glaziophyton Franch. than it is to other 

Aulonemia species.

KEY TO THE SPECIES OF AULONEMIA

1a. Sheaths without marginal fimbriae; lemmas 6.5–8 mm long, awns absent . . . . . . . . . . . . . . . . . . . . . . . . . . . A. trianae

1b. Sheaths with marginal fimbriae along their entire length; lemmas 8–12.5 mm long, awns 1.5–5 mm long or absent . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. ximenae

Aulonemia trianae (Munro) McClure, Smithsonian Contr. Bot. 

9: 61. 1973. Arundinaria trianae Munro, Trans. Linn. Soc. 

London 26(1): 25. 1868.

Small bamboos. Culms erect or scandent, 0.5–2.5(–6) m tall, 

5–10 mm in diameter; internodes 3–25 cm long. Culm leaves: deciduous; sheaths 5–10 cm long, glabrous, apices sparsely fimbriate, margins not fimbriate; blades 4–11.5 × 0.6–2.2 cm, reflexed. 

Foliage leaves: sheaths glabrous, apices sparingly fimbriate, 

fimbriae ~1 cm long, margins not fimbriate; blades lanceolate, 

5–15 × 1–2.5 cm, reflexed, apices acuminate to slightly navicular, 

bases rounded, glabrous above, glabrous or scabrid below. Panicles 10–13 cm long. Spikelets 1–1.8 × 0.3–0.5 cm, with 5–6(–8) 

fertile florets; glumes acuminate to shortly subulate, awnless; 

lemmas 6.5–8 mm long, awns absent.

Found in South America (Colombia and Venezuela). In Venezuela, found only in the Andes (Lara, Mérida, Táchira, and Trujillo; Map 263). Páramo del Pumar; 2,500–2,600 m.

This species forms dense colonies.

MAP 262. Arthrostylidium venezuelae occurrence in Venezuela.
218 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Aulonemia ximenae L. G. Clark et al., Bamboo Sci. Cult. 20(1): 

3, fig. 1. 2007.

Small bamboos. Culms erect, climbing or scandent, to 2(–5) 

m tall, 3–6(–10) mm in diameter; internodes 10–26 cm long. Culm 

leaves: tardily deciduous; sheaths 10–18 cm long, sparsely hispid 

or glabrous, marginal fimbriae present the full length of sheath, 

apical fimbriae 7–15 mm long; blades lanceolate, 3–10 × 0.8–2.5 

cm, strongly reflexed. Foliage leaves: sheaths glabrous below, 

glabrous to puberulent above, marginal fimbriae present the full 

length of sheath, apical fimbriae 7–15 mm long; blades lanceolate, 

8–15(–17) × 1–2.5(–2.8) cm, reflexed, bases truncate, ± oblique, 

apices acuminate to slightly navicular, glabrous above, glabrous to 

puberulent below. Panicles 15–20 cm long. Spikelets 1.3–3(–3.6) 

× 0.3–0.6 cm, with (4–)5–9 fertile florets; glumes acute to awned; 

lemmas 8–12.5 mm long, awns 1.5–5 mm long or absent.

Found in South America (Colombia and Venezuela). In Venezuela, found only in the Andes (Lara, Mérida, Táchira, and Trujillo; Map 264). Páramo del Pumar; 2,500–2,600 m.

Axonopus P. Beauv.

Axonopus P. Beauv., Ess. Agrostogr. 12, 154. 1812.

Centrochloa Swallen, J. Wash. Acad. Sci. 25: 192. 1935.

Ophiochloa Filg. et al., Novon 3: 360. 1993.

Perennial, rarely annual plants; caespitose, stoloniferous 

or rhizomatous. Culms herbaceous, often decumbent, erect to 

ascending. Sheaths keeled, open; ligules membranous, ciliate; 

blades flat, folded or involute, linear to linear-lanceolate. Inflorescences terminal or axillary panicles, with 2 to many spikelike, 

often digitate branches; branches 3-angled; spikelets solitary, in 

2 rows, appressed to rachises; disarticulation below the glumes. 

Spikelets subsessile or sessile, compressed dorsally, with 2 florets; 

lower florets sterile or staminate; upper florets bisexual. Lower 

glumes absent; upper glumes and lemmas subequal, membranous, 2–5-nerved; lower paleas absent; upper lemmas rigid, usually glabrous or with an apical tuft of hairs; upper paleas similar 

in texture to upper lemmas; anthers 3; styles 2, separate. Caryopses ellipsoid to ovoid, compressed. [Panicoideae.]

Approximately 85 species native to tropical and subtropical North America (USA), Mexico, Central America, the West 

Indies, and South America (Colombia, Venezuela, Trinidad and 

Tobago, the Guianas, Ecuador, Peru, Bolivia, Brazil, Chile, Paraguay, Argentina, and Uruguay); various species introduced and 

naturalized in Europe, Africa, Asia, Australia, and the Pacific 

Islands. Thirty-four species occur in Venezuela.

References. Crins (1991); Giraldo-Cañas (2007, 

2008); López and Morrone (2012); Morrone et al. (2012); Webster (1988); Webster et al. (1989).

Spikelet orientation is the only morphological character that 

separates Axonopus from the closely related Paspalum L. In the 

former genus fertile lemmas and upper glumes are turned away 

from the inflorescence rachis.

Axonopus compressus (Sw.) P. Beauv., Ess. Agrostogr. 12, 154, 

167. 1812; Davidse, in Berry et al., Fl. Venez. Guayana 8: 

50–51. 2004; Briceño, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 2: 574, fig. 28. 2010. Millium compressum

Sw., Prodr. 24. 1788.

Perennial plants. Culms 10–50(–85) cm tall; nodes glabrous 

or pubescent. Leaves basal and cauline; sheaths compressed, 

pubescent; ligules 0.2–0.5 mm long; blades flat, subulate, 

linear-lanceolate to lanceolate, 5–20 × 0.5–1 cm, apices often 

ciliate, bases ciliate, glabrous or sparingly pubescent. Inflorescences terminal and axillary, composed of 1–5(–6) digitate 

branches; branches 3–10(–15) cm long; rachises 3-angled, 

MAP 264. Aulonemia ximenae occurrence in Venezuela.

MAP 263. Aulonemia trianae occurrence in Venezuela.
NUMBER 100 • 219

glabrous. Spikelets subsessile, 2–2.5 mm long, ovoid, ellipsoid 

or lanceoloid, pubescent; upper glumes and lower lemmas equal, 

2–4(–5)-nerved, veins pubescent; upper lemmas and paleas 1.5–2 

mm long, ellipsoid. Caryopses ellipsoid to ovoid, ~1 mm long.

Found in North America (USA), Mexico, Central America, 

the West Indies, South America (all upper-level political units), and 

naturalized in tropical and subtropical regions of the Old World. 

Widely distributed in Venezuela (Amazonas, Anzoátegui, Apure, 

Aragua, Barinas, Bolívar, Carabobo, Cojedes, Delta Amacuro, 

Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, 

and Zulia; Map 265). In the park, found in wet, disturbed open 

areas on both slopes of Guaramacal; 1,800–2,000 m.

This species is used as a lawn and forage grass. It is, however, weedy and often found in wet disturbed areas.

Bromus L.

Bromus L., Sp. Pl. 76. 1753, nom. cons.

Annual, biennial or perennial plants; caespitose or rarely 

rhizomatous. Culms unbranched. Sheaths tubular, closed; blades 

linear, flat. Inflorescences terminal, solitary panicles, lax or contracted. Spikelets with 1 to many fertile florets, laterally compressed; disarticulation between the glumes and between the 

florets. Glumes 2, herbaceous, unequal, acute, smaller than florets, 

1–9-nerved; lemmas herbaceous or subcoriaceous, 5–9-nerved, 

rounded or keeled, mucronate or awned; paleas 2-nerved, carinate, smaller than lemmas; lodicules 2, membranous; anthers 3; 

styles 2. Caryopses linear or ellipsoid. [Pooideae.]

A cosmopolitan genus of 140–160 species found mostly in temperate regions and at higher elevations in the tropics. In the Americas, found in North America (Canada and USA), Mexico, Central 

America, and South America (Colombia, Venezuela, Ecuador, Peru, 

Bolivia, Brazil, Chile, Paraguay, Argentina, and Uruguay); exotic in 

the West Indies. Many species are Andean; 4 or 5 occur in Venezuela.

References. Peterson and Planchuelo (1998); 

Pinto-Escobar (1981); Planchuelo and Peterson (2000); Saarela 

et al. (2007); Tucker (1996).

Bromus catharticus Vahl, Symb. Bot. 2: 22. 1791; Briceño, in 

Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 579, 

fig. 32. 2010.

Material from Venezuela and Guaramacal can be referred to 

the nominate variety.

Bromus catharticus var. catharticus

Annual or biennial, occasionally perennial plants. Culms erect 

or somewhat decumbent, 25–100(–120) cm tall. Leaf sheaths glabrous or sparingly pubescent; ligules 2–5 mm long; blades 10–20 

× 0.2–0.6 cm, glabrous or sparingly pubescent. Panicles 10–30 

cm long, 5–9-flowered, erect or lax. Spikelets 15–25 mm long, 

4–11-flowered. Glumes ovate, glabrous; lower glumes 6–12 mm 

long, 3–7-nerved; upper glumes, 8–14 mm long, (5–)7–9-nerved; 

lemmas ovate, 10–15 mm long, 7–11-nerved, glabrous or minutely pubescent, apices acute, aristate, awns 0.5–4(–5) mm long; 

paleas 5–10 mm long, strongly keeled, ciliate. Caryopses sulcate.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, Bolivia, Brazil, Chile, Paraguay, Argentina, and Uruguay); introduced in North America (Canada and USA), Mexico, Central 

America, the West Indies, Trinidad and Tobago, Europe, Africa, 

Asia, Australia, and the Pacific Islands. In Venezuela, reported from 

the Andes (Lara, Mérida, Táchira, and Trujillo) and the Cordillera 

de la Costa (Distrito Federal and Miranda; Map 266). Our sole 

record is from open areas near the Laguna de los Cedros; 2,000 m.

MAP 265. Axonopus compressus occurrence in Venezuela.

MAP 266. Bromus catharticus var. catharticus occurrence in 

Venezuela.
220 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Bromus catharticus is highly variable and ecologically diverse. Nonetheless, in South America three varieties currently 

are recognized and can be separated by the length of their lemma 

awns. Bromus catharticus var. rupestris (Speg.) Planchuelo & 

P. M. Peterson, restricted to Argentina, has mucronate lemmas or 

lemmas with awns 0.3–0.5(–1) mm long, whereas B. catharticus

var. elatus (É. Desv.) Planchuelo (=B. cebadilla Steud.), found in 

Peru, Chile, and Argentina, has lemmas with awns 5–10 mm long.

Bromus catharticus is used as a forage grass, which explains 

in part its extensive nonnative range.

Calamagrostis Adans.

Calamagrostis Adans., Fam. Pl. 2: 31, 530. 1763.

Deyeuxia Clarion ex P. Beauv., Ess. Agrostogr. 43. 1812. Calamagrostis sect. 

Deyeuxia (Clarion ex P. Beauv.) Dumort., Observ. Gramin. Belg. 126. 

1823 [1824].

Perennial plants; caespitose, rhizomatous or not, rarely stoloniferous. Culms unbranched, erect, ascending or decumbent. 

Leaves cauline, few; sheaths open, glabrous or scabrous; ligules membranous, glabrous or pilose; blades filiform or linear, 

plane, convolute or conduplicate. Inflorescences terminal panicles, narrow or contracted, dense or lax. Spikelets lanceolate, 

1(2)-flowered, somewhat laterally compressed; disarticulation 

above the glumes. Rachilla briefly prolonged or not. Glumes 2, 

subequal, lanceolate, 1–3-nerved, acute or acuminate, glabrous 

or scabridulous along midnerve; lemmas ellipsoid, 3(–5)-nerved, 

glabrous or scabrous, almost always awned; awn arising from 

near the base to near the apex, straight or geniculate, exserted 

(our species) or not; callus bearded or not; paleas membranous, 

± equal to lemmas in length. Lodicules 2. Stamens (1–)3. Ovary 

glabrous. Styles separate, very short. Stigmas 2, plumose. Caryopses with adherent pericarp, usually glabrous. [Pooideae.]

A genus of ~270 species found worldwide in temperate 

areas and at higher elevations in the tropics. In the Americas, 

known to occur in North America (Greenland, Canada, and 

USA), Mexico, Central America, the Greater Antilles (Hispaniola), and South America (Colombia, Venezuela, Ecuador, Peru, 

Bolivia, Brazil, Chile, Paraguay, Argentina, and Uruguay). At 

least 16 species occur in Venezuela.

References. Phillips and Chen (2003); Rúgolo de 

Agrasar (1978); Saarela et al. (2010); Tucker (1996); Wasiljew (1960).

The genera Calamagrostis, Deyeuxia, and Agrostis are closely 

related and form an intergrading complex of incompletely separated entities (Phillips and Chen, 2003). Although Agrostis is always maintained separately, Calamagrostis and Deyeuxia are either 

considered to be separate genera (Rúgolo de Agrasar, 1978) or Deyeuxia is synonymized under Calamagrostis (Soreng and Greene, 

2003), which is the treatment adopted here (see also the key to genera of Poaceae). Phillips and Chen (2003) argue that recognition of 

only two genera in this complex is not a tenable taxonomic solution as the boundaries between Agrostis and both Calamagrostis

and Deyeuxia are as ill defined as those between Calamagrostis and 

Deyeuxia. Inasmuch as morphology does not resolve the problem, 

a satisfactory taxonomic treatment of this generic complex awaits a 

molecular phylogenetic study of the entire group worldwide.

KEY TO THE SPECIES OF CALAMAGROSTIS

1a. Awn slightly exceeding lemma in length by <2 mm; hairs on callus scarcely visible or absent . . . . . . . . . . C. bogotensis

1b. Awn exceeding lemma in length by >2 mm; hairs on callus visible . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Culms ± erect; leaf blades linear, 0.2–0.4 cm in width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. planifolia

2b. Culms decumbent or stoloniferous; leaf blades linear-lanceolate, 0.5–0.8 cm in width . . . . . . . . . . . . . . . C. sp. A.

Calamagrostis bogotensis (Pilg.) Pilg., Bot. Jahrb. Syst. 42: 60. 

1908; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 584. 2010. Deyeuxia bogotensis Pilg., Bot. 

Jahrb. Syst. 25: 712. 1898.

Deyeuxia nuda Pilg., Bot. Jahrb. Syst. 27: 29. 1899. Calamagrostis nuda

(Pilg.) Pilg., Bot. Jahrb. Syst. 42: 60. 1908.

Rhizomatous. Culms 30–140 cm tall, ± erect. Sheaths glabrous; ligules eciliate; blades linear, convolute, 10–40 × 0.2–0.5 

cm, glabrous or minutely scabrous above and below. Inflorescences compact, 5–22 × 1–2.5 cm; spikelets solitary, 3.5–4.5 mm 

long. Glumes subequal, 3.5–4.5 mm long, acute or acuminate, 

1(–3)-nerved, glabrous or scabrous; lemmas 2.5–3.5 mm long, 

5-nerved, apices bifid or 4-toothed, awned; awn ~4 mm long, dorsal, inserted near base of lemma and slightly exceeding (<2 mm) it 

in length; callus hairs scarcely visible or absent. Stamens (1) 3.

MAP 267. (Right) Calamagrostis bogotensis occurrence in Venezuela.
NUMBER 100 • 221

Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, and Ecuador). In Venezuela, found only in the Andes (Lara, Mérida, Táchira, and Trujillo; 

Map 267). In the park, found in the Páramo de Guaramacal, 

Páramo del Pumar, and Páramo de Vicuyal; 2,600–3,150 m.

Calamagrostis planifolia (Kunth) Trin. ex Steud., Nomencl. 

Bot., 2nd ed., 1: 251. 1840; Briceño, in Morillo et al., Bot. 

Ecol. Monocot. Páramos Venez. 2: 592, fig. 40. 2010. Deyeuxia planifolia Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 

1:117. 1815 [1816]; ibid. [qu.] 1: 145. 1815 [1816].

Calamagrostis pittieri auct., non Hack.; Dorr et al., Contr. U.S. Natl. Herb. 

40: 56. 2000 [2001].

Rhizomatous. Culms 20–30(–100) cm tall, ± erect. Sheaths 

glabrous, scabrous or pilose; ligules ciliolate; blades linear, flat, 

5–12 × 0.2–0.4 cm, scabrous above and below. Inflorescences 

lax, 8–15 × 1.5–4 cm; spikelets solitary, 3.8–5 mm long. Glumes 

equal or subequal, 3–5 mm long, acute or acuminate, 1–3-nerved, 

smooth or asperulous, lateral nerve scabrous; lemmas 3.6–5 mm 

long, 5-nerved, apices bifid, awned; awn 5–7 mm long, dorsal, 

inserted above the middle of the lemma and exceeding (>2 mm) 

it in length; callus pilose. Stamens 2.

Found in Central America (Costa Rica) and South America (Colombia, Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, found at higher elevations in the Andes (Lara, Mérida, 

Táchira, and Trujillo), the Sierra de Perijá (Zulia), and the 

Cordillera de la Costa (Distrito Federal; Map 268). A subpáramo and páramo species found in the park in the Páramo de 

Guaramacal, Páramo del Pumar, and Fila de Agua Fría; (2,500–)

2,600–3,100 m.

The inflorescences of this species are often purplish. Calamagrostis planifolia is very similar to C. pittieri Hack., and the 

two species are sometimes considered synonymous (Soreng and 

Greene, 2003). Zuloaga et al. in Hokche et al. (2008), however, 

recognized both species in the flora of Venezuela and gave them 

more or less overlapping ranges. Costa Rican floras (Pohl, 1980; 

Morales in Hammel et al., 2003b) also accept C. pittieri but consider it to be endemic to Costa Rica.

Calamagrostis sp. A

Calamagrostis chaseae auct., non Luces; Dorr et al., Contr. U.S. Natl. Herb. 

40: 56. 2000 [2001].

Stoloniferous. Culms 0.8–1 m tall, decumbent or stoloniferous. Sheaths glabrous; ligules eciliate; blades linear-lanceolate, flat, 

15–25 × 0.5–0.8 cm, glabrous or slightly scabrous. Inflorescences 

lax, 10–20 × 3–5 cm; spikelets solitary, 3.5–4 mm long. Glumes 

equal, 3–4.4 mm long, 1-nerved, glabrous or scabrous; lemmas 

3–4.2 mm long, 5-nerved, apices bifid, awned; awn 3.5–6 mm 

long, dorsal, inserted above the middle of the lemma and exceeding (>2 mm) it in length; callus puberulent. Stamens 3(2).

Evidently restricted to the Andes of Venezuela (Trujillo; 

Map 269). At present only known from the park, where it occurs in the Páramo de Guaramacal and on both slopes below the 

summit; (2,500–)2,700–3,000 m.

This taxon is similar to the Andean Calamagrostis effusa

(Kunth) Steud. but differs in having decumbent or stoloniferous 

culms and broad, flat leaf blades.

Cenchrus L.

Cenchrus L., Sp. Pl. 1049. 1753.

Pennisetum Rich., in Persoon, Syn. Pl. 1: 72. 1805; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 58. 2000 [2001].

Annual or perennial plants; caespitose, rhizomatous or 

stoloniferous. Culms erect, decumbent, geniculate or creeping, 

herbaceous, often branched; internodes solid or hollow. Sheaths 

MAP 268. Calamagrostis planifolia occurrence in Venezuela. MAP 269. Calamagrostis sp. A occurrence in Venezuela.
222 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

open; ligules membranous or ciliate; blades flat or folded, sometimes pseudopetiolate. Inflorescences terminal, terminal and axillary, or only axillary spicate panicles, with fascicles (i.e., highly 

reduced branches); rachises terete, with (1–)5 to many fascicles; 

fascicle axes with (1–)3–100+ bristles and 1–12 spikelets. Bristles 

free or fused at base; disarticulation at base of fascicles, sometimes also beneath upper florets. Spikelets with 2 florets. Lower 

glumes ovoid, rudimentary or absent, 0–5-nerved; upper glumes 

equal or subequal to spikelets, 0–11-nerved; lower florets sterile 

or staminate; lower lemmas as long as spikelets, membranous, 

3–15-nerved; lower paleas shorter than or equaling lemmas; 

upper lemmas membranous to coriaceous, 5–12-nerved; upper 

paleas shorter than or subequal to lemmas; lodicules 0 or 2, glabrous; anthers (0) 3. [Panicoideae.]

A genus of 100–160 species found in tropical and subtropical regions worldwide, including North America (Canada and 

USA), Mexico, Central America, the West Indies, and South 

America (all country-level political units). Fourteen species occur 

in Venezuela.

References. Chase (1920, 1921); Chemisquy et al. 

(2010); Crins (1991); Morrone et al. (2012); Türpe (1983); Webster (1988); Webster et al. (1989).

KEY TO THE SPECIES OF CENCHRUS

1a. Low creeping plants; inflorescences axillary, partially or completely hidden in the upper leaf sheaths at maturity (only 

stamens and stigmas exserted) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. clandestinus

1b. Erect or decumbent plants; inflorescences terminal or terminal and axillary, fully exserted at maturity . . . . . . . . . . . . 2

2a. Panicles terminal and axillary, 5–10 cm long; bristles ± flexuous; leaf blades 10–25 × 1–2 cm . . . . . C. bambusiformis

2b. Panicles terminal on the primary culm and leafy branches only, 10–25 cm long; bristles soft and silky; leaf blades 

10–40 × 2–4 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. peruvianus

Cenchrus bambusiformis (E. Fourn.) Morrone, in Chemisquy et 

al., Ann. Bot. (Oxford), n.s., 106: 127. 2010. Gymnotrix 

bambusiformis E. Fourn., Mexic. Pl. 2: 48. 1886 (“Gymnothrix”). Pennisetum bambusiforme (E. Fourn.) Hemsl. ex B. 

D. Jacks., Index Kew. 2: 458. 1895; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 58. 2000 [2001].

Perennial plants; caespitose. Culms erect or decumbent, 

1–2(–4) m × 3–7 mm, branched; internodes hollow. Sheaths glabrous, papillose-ciliate on upper margins and auricles; ligules 

membranous and ciliate; blades lanceolate, 10–25 × 1–2 cm, flat, 

apices acuminate, ± hirsute above and below. Inflorescences axillary and terminal panicles; fascicles 5–10 × 1.5–2.5 cm; rachises 

scabrous. Bristles numerous, unequal, 5–15 mm long. Spikelets 

ovate, 3.5–4.5 mm long. Glumes unequal; lower glumes very 

small; upper glumes about 1⁄2 the length of spikelets; lower lemmas longer than upper lemmas; lodicules 2; anthers 3.

Found in Mexico, Central America, and South America (Colombia, Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, 

collected in the Andes (Barinas, Lara, Mérida, Portuguesa, 

Táchira, and Trujillo) and in the Cordillera de la Costa (Aragua 

and Distrito Federal; Map 270). In the park, known from both 

slopes of Guaramacal; 1,600–2,100 m.

• Cenchrus clandestinus (Hochst. ex Chiov.) Morrone, in 

Chemisquy et al., Ann. Bot. (Oxford), n.s., 106: 127. 2010. 

Pennisetum clandestinum Hochst. ex Chiov., Annuario 

Reale Ist. Bot. Roma 8: 41, t. 5, fig. 2. 1903, nom. cons. 

prop.; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 659, fig. 92. 2010.

Pennisetum longistylum Hochst. ex A. Rich., Tent. Fl. Abyss. 2: 388. 1850, 

nom. rej. prop.

Perennial plants; rhizomatous or stoloniferous. Culms 

prostrate, 3–15(–45) cm long, branched. Sheaths keeled, pilosepubescent, somewhat inflated; ligules ciliate; blades flat or conduplicate, linear, 6–10 × 0.3–0.4 cm, apex obtuse, glabrous 

above and below or with scattered long hairs. Inflorescences 

compound panicles, reduced to a few (1–6) subsessile spikelets 

hidden in the upper sheaths, with only the stamens and stigmas 

exserted. Bristles 3–15 in principal whorl, unequal, shorter than 

the spikelets. Spikelets narrowly lanceolate, 10–20 mm long. 

Glumes absent or obscure; lower glumes reduced to brief scales; 

upper glumes 1–3 mm long; lower and upper lemmas equal in 

length; lodicules 0; anthers 3.

MAP 270. Cenchrus bambusiformis occurrence in Venezuela.
NUMBER 100 • 223

Native to Africa; introduced and naturalized throughout the 

tropics and subtropics. In the Americas, found in North America 

(USA), Mexico, Central America, the West Indies, and South 

America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, 

Chile, Paraguay, Argentina, and Uruguay). In Venezuela, found 

only in the Andes (Lara, Mérida, Táchira, and Trujillo; Map 

271). In the park, encountered in wet pastures on the north slope 

of Guaramacal; 2,100 m.

This species, well known in South America as “kikuyo” or 

“pasto kikuyo,” may have been introduced to the Americas as late 

as the 1920s, probably through agricultural experiment station 

activity (Parsons, 1972) as opposed to accident. In South America 

it prefers higher elevations, which stands in contrast to other African forage grass introductions that favor lower, hotter elevations.

Gutiérrez (2014) proposed that the name Pennisetum clandestinum Hochst. ex Chiov. be conserved over the earlier P. 

longistylum Hochst. ex A. Rich. If his proposal fails, then the 

correct name for this species will be a combination based on the 

latter species name.

Cenchrus peruvianus (Trin.) Morrone, in Chemisquy et al., Ann. 

Bot. (Oxford), n.s., 106: 128. 2010. Pennisetum peruvianum Trin., Linnaea 10: 295. 1836; Ortega et al., BioLlania 

5: 42. 1987; Dorr et al., Contr. U.S. Natl. Herb. 40: 58. 

2000 [2001].

Perennial plants. Culms erect or decumbent, 1–3 m × 4–6 

mm, branched; internodes hollow. Sheaths glabrous; ligules 

ciliate; blades lanceolate, 10–40 × 2–4 cm, flat, apices longacuminate, glabrous above, pubescent below. Inflorescences 

terminal panicles (on primary culm and leafy branches only); fascicles 15–25 × 3–4.5 cm; rachises scabrous. Bristles numerous, 

± equal, very dense and silky, 2–3 cm long. Spikelets lanceolate, 

3–4 mm long. Glumes unequal; lower glumes much smaller than 

spikelets; upper glumes ± equal in length to spikelets. Lower and 

upper lemmas ± equal in length; lodicules 0; anthers 3.

Found in South America (Colombia, Venezuela, Ecuador, 

and Peru); adventive in the Pacific Islands. In Venezuela, found 

only in the Andes (Lara, Mérida, Táchira, and Trujillo; Map 

272). In the park, known from both slopes of Guaramacal; 

1,500–1,800(–2,700) m.

We also have seen a fragmentary specimen (Linden 329, US) 

of this species consisting of a few florets that is labeled “Caracas,” but we are not convinced that Cenchrus peruvianus occurs 

in the Cordillera de la Costa as this incomplete specimen is the 

only record from a relatively well-collected area.

Chusquea Kunth

Chusquea Kunth, J. Phys. Chim. Hist. Nat. Arts 95: 151. 1822.

Neurolepis Meisn., Pl. Vasc. Gen. 1: 426. 1843; ibid. 2: 325. 1843; Dorr et 

al., Contr. U.S. Natl. Herb. 40: 57. 2000 [2001].

Swallenochloa McClure, Smithsonian Contr. Bot. 9: 106. 1973.

Woody bamboos; caespitose; often forming colonies. Rhizomes absent or present; pachyform, leptomorph or both pachyform and leptomorph. Culms unarmed or armed by a ring of 

short, sharp aborted roots at aboveground nodes (C. fendleri

Munro only), erect or arched, sometimes scandent or climbing, 

solid, hollow or solid and becoming fistulose; internodes subequal, cylindric or terete; aerial culms branched or not. Branching extravaginal, infravaginal or intervaginal. Culm leaves: 

sheaths persistent or deciduous; blades triangular or lanceolate 

to linear. Foliage leaves: sheaths erect, without auricles or oral 

MAP 271. Cenchrus clandestinus occurrence in Venezuela. MAP 272. Cenchrus peruvianus occurrence in Venezuela.
224 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

setae; ligules external and internal or internal only, membranous; blades lanceolate to elliptic or oblong, basal or not, midrib 

prominent or not, articulate, persistent or deciduous; pseudopetiolate or sessile. Inflorescences axillary or terminal panicles, 

rarely racemes, open or contracted. Spikelets terete or laterally 

compressed, pedicellate; consisting of 2(4–5) glumes, 2 sterile 

lemmas, and 1 bisexual floret, rachilla not prolonged above the 

terminal fertile floret; disarticulation above the glumes and below 

the sterile lemmas. Lodicules (2) 3. Stamens (2) 3(–6). Ovary glabrous. Stigmas 2. Caryopses rarely seen. [Bambusoideae.]

A neotropical genus of 135–160(–200) species found in 

Mexico, Central America, the Greater Antilles, and South 

America (Colombia, Venezuela, Trinidad and Tobago, Guyana, 

Ecuador, Peru, Bolivia, Brazil, Chile, Paraguay, Argentina, and 

Uruguay). Twenty-two species occur in mountainous areas of 

Venezuela.

References. Calderón and Soderstrom (1980); 

Clark (1989, 1997); Clark and Ely (2011, 2013); Fisher et al. 

(2009); Judziewicz et al. (1999); McClure (1973); Niño et al. 

(2006); Soderstrom (1969).

KEY TO THE SPECIES OF CHUSQUEA

1a. Aerial culms abundantly branching at the middle and upper nodes; foliar leaf blades <40 cm long . . . . . . . . . . . . . . . 2

2a. Plants scandent, stoloniferous; foliage leaf blades ovate to narrowly ovate, 2–5 cm wide . . . . . . . . . . . . C. serpens

2b. Plants erect, arching or scandent, never stoloniferous; foliage leaf blades lanceolate or linear to linear-lanceolate, 

0.2–2 cm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Foliage leaf blades lanceolate (rarely linear-lanceolate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Basal nodes of culms usually with curved adventitious roots; bases of foliage leaf blades attenuate or rounded, 

apices attenuate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. fendleri

 4b. Basal nodes of culms without adventitious roots; bases of foliage leaf blades rounded or truncate, apices 

acuminate-subulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. tessellata

 3b. Foliage leaf blades linear to linear-lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 5a. Culms scandent, long arching; 6–8 m long, 1–3 cm in diameter . . . . . . . . . . . . . . . . . . . . . . C. multiramea

 5b. Culms erect, sometimes culm and branch tips pendulous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 6a. Culms completely erect, 0.1–1.5(–3) m long, 0.3–1 cm in diameter; culm leaves smooth to scabrous or 

often pubescent near base; spikelets pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. angustifolia

 6b. Culms erect, but culm and branch tips pendulous, 2–6 m long, 0.5–4 cm in diameter; culm leaves hispid 

below; spikelets glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. spencei

1b. Aerial culms never branched; foliar leaf blades (70–)150–300 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

7a. Panicle axis and branches glabrous; pedicels longer than spikelets . . . . . . . . . . . . . . . . . . . . . . . . . . . C. spectabilis

7b. Panicle axis and branches densely and soft pubescent; pedicels shorter than spikelets . . . . . . . . . . . . . . . . C. mollis

Chusquea angustifolia (Soderstr. & C. E. Calderón) L. G. Clark, 

Ann. Missouri Bot. Gard. 74: 428. 1987; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 598–599, 

fig. 45, foto 7. 2010. Swallenochloa angustifolia Soderstr. & 

C. E. Calderón, Brittonia 30: 303. 1978.

Shrubby, woody bamboos; forming dense colonies. Culms 

0.1–1(–3) m × 3–12 mm, erect; internodes 4–15 cm long, solid, 

becoming fistulose, yellowish tan, smooth, waxy. Culm leaves: 

sheaths ± rectangular, 3–6(–10) cm long, smooth to scabrous or 

pubescent at base; blades narrowly triangular, 1–3(–5) cm long, 

erect, sparingly pubescent above and below, persistent. Branching intravaginal, up to 9 branches present in mature branch complement. Foliage leaves: 4–8(–13) per complement; sheaths 4–10 

× 3–6 cm, usually glabrous, persistent; blades linear to linearlanceolate, 4–11 cm × 3–7 mm, erect, bases attenuate, margins 

minutely scabrous, apices setose-subulate, glabrous above, usually pilose below; pseudopetiolate. Panicles narrow, congested, 

3–9(–14) cm long, green to purple; primary panicle branches appressed, 1–2 cm long. Spikelets obovate-lanceolate, 4.5–7.4 mm 

long, pubescent.

Found in South America (Colombia and Venezuela). In 

Venezuela, found in the Andes (Apure, Lara, Mérida, Táchira, 

and Trujillo) and in the Sierra de Perijá (Zulia; Map 273). This 

species occurs in subpáramo and páramo throughout the park; 

(2,500–)2,700–3,100 m.

Chusquea angustifolia is closely related to C. spencei Ernst, 

and the two species are difficult to distinguish (see key). The 

latter species is much larger in stature than the former, and as 

detailed by Clark (1989), C. spencei has abaxially hirsute, glabrescent culm leaves; more complex branch complements; longer, narrower foliage leaves; somewhat longer inflorescences; 

and glabrous spikelets.

In Guaramacal, Chusquea angustifolia is variable in stature. Plants in exposed locations are more compact than those 

shielded by rocks or other barriers that provide shelter from the 

prevailing winds.

Chusquea angustifolia is reported to occur in Sucre, Venezuela (Clark and Ely, 2011), but we have not seen the voucher, and 

this record needs to be confirmed, especially as the closely related 

C. spencei is known to occur in the Cordillera de la Costa.
NUMBER 100 • 225

Chusquea fendleri Munro, Trans. Linn. Soc. London 26(1): 61. 

1868.

 FIGURE 26A

Woody bamboos. Culms 4–8 m × 10–20 mm, arching or 

scandent; internodes solid, 4–35 × 2–3 cm, reddish-purple, scabrous; nodes with recurved adventitious roots that often pierce 

the bases of culm leaves. Branch complement: many clumped, 

subequal branches, 1–3 mm in diameter (thinner than stems). 

Culm leaves: sheaths triangular, pubescent to scabrous below, 

persistent; blades 4–6.5(–15) cm long, margins smooth. Foliage 

leaves: 4–6 per complement; sheath apices conspicuously longciliate, surface glabrous; blades lanceolate (rarely narrowly lanceolate), 3–15 cm × (5–)10–20 mm, bases attenuate or rounded, 

margins ciliate, apices attenuate, glabrous above, pubescent 

below or not; pseudopetiolate. Panicles spiciform, linear, 8–12

(–20) cm long. Spikelets elliptic, 4–10 mm long.

Found in South America (Colombia and Venezuela). In Venezuela, recorded from the Andes (Lara, Mérida, and Trujillo) 

and the Cordillera de la Costa (Anzoátegui, Aragua, and Distrito Federal; Map 274). Found in cloud forest on both slopes 

of Guaramacal and in shrubby páramo along the main ridge of 

the park (i.e., Fila de Los Recostaderos–Páramo de Guaramacal–Páramo del Pumar); 1,800–3,100 m.

As noted in the generic description, this is the only “armed” 

species of Chusquea, with its short, sharp aborted roots piercing 

the bases of culm leaves at nodes.

Chusquea mollis (Swallen) L. G. Clark, in Fisher et al., Syst. 

Bot. 34: 681. 2009. Neurolepis mollis Swallen, J. Wash. 

Acad. Sci. 21: 14. 1931; Dorr et al., Contr. U.S. Natl. Herb. 

40: 57. 2000 [2001]; Briceño, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 2: 646, fig. 83, foto 11 (as Neurolepis glomerata). 2010, excluding fotos 12–14.

Neurolepis glomerata auct., non Swallen; Dorr et al., Contr. U.S. Natl. Herb. 

40: 57. 2000 [2001].

Woody bamboos; forming dense colonies. Culms 1.5–4.5 

m tall, erect, unbranched above the base. Sheaths longer than 

adjacent culm internodes, smooth, glabrous above and below, setose at the mouth; ligules membranous, ± lacerate above, 1–2 cm 

long; blades lanceolate, 1.5–2 m × 3–5.5 cm, bases attenuate, 

margins minutely scabrous, apices acuminate, sparsely pilose 

to glabrescent above, glabrous below; pseudopetiolate. Panicles lanceolate to ovate, narrow to expanded, 0.6–1.5 m long, 

contracted about primary branches (immature inflorescences?), 

dark purple to blackish; primary branches appressed or ascending, 5–15 cm long, bearing spikelets almost to the base; panicle 

axis and branches densely and soft pubescent. Spikelets ovate, 

2–4 mm long.

Found in South America (Colombia and Venezuela). In 

Venezuela, evidently restricted to the Andes (Mérida, Táchira, 

and Trujillo; Map 275). In the park, found in cloud forest on 

the north slope and exposed areas of subpáramo and páramo; 

(2,500–)3,000–3,100 m.

Dorr et al. (2000) recognized two species in this complex 

as occurring in Guaramacal, but reexamination of the available 

collections leads us to believe that there is only one species collected at different stages of flowering. The inflorescences on most 

of the fertile collections from Guaramacal are either immature 

or not fully expanded and appear to be narrow, lanceolate or 

spikelike panicles. The inflorescence on one collection (Dorr et 

al. 7404), however, appears to be fully expanded, the primary 

MAP 273. Chusquea angustifolia occurrence in Venezuela.

MAP 274. Chusquea fendleri occurrence in Venezuela.
226 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

panicle branches are ascending, and the inflorescence is ± ovate. 

Spikelets in both inflorescence forms appear to be identical. None 

of these Guaramacal collections is a good match for Chusquea 

glomerata (Swallen) Dorr, which was described from the Venezuelan Guayana. Chusquea glomerata has a well-developed pulvinus 

at the base of each panicle branch, but our Guaramacal material 

has inconspicuous and scarcely developed pulvini. Our material 

best matches the Andean C. mollis, which has spreading panicle 

branches and puberulent sterile lemmas. The inflorescence on the 

type specimen of C. mollis is immature and matches the narrow, 

lanceolate or spikelike panicles in our immature material. A third 

member of this complex, C. magnifolia L. G. Clark described from 

the Cordillera de la Costa, differs from C. mollis in that the sterile 

lemmas are not short-pubescent toward the summit. Clearly, all 

three of these species deserve closer taxonomic scrutiny.

In Guaramacal it is almost impossible to distinguish sterile 

Chusquea mollis from sterile C. spectabilis; foliage leaf blades 

are generally narrower in the former (3–5.5 vs. 4.5–15 cm wide), 

and sometimes hairs can be seen on the upper foliage leaf surface 

of the former (the surface of the latter is always glabrous).

Briceño in Morillo et al. (2010) confounded fertile material 

of Chusquea mollis (cited as Neurolepis mollis) with C. spectabilis; the photographs (fotos 12–14) that they published of 

spikelets and inflorescences of C. mollis are, in fact, images of 

C. spectabilis.

Chusquea multiramea L. G. Clark & F. Ely, Syst. Bot. 38: 1091, 

fig. 1D–H. 2013.

Chusquea purdieana auct., non Munro; Dorr et al., Contr. U.S. Natl. Herb. 

40: 57. 2000 [2001].

Woody bamboos; forming colonies. Culms 4–8 m × 1–3 cm, 

scandent, long arching; internodes 10–30 cm long, solid; nodes 

without adventitious roots. Branch complement: many (100–

150), dense subequal branches, 0.5–1 mm in diameter (thinner 

than stems). Culm leaves: sheaths persistent, bright brown at the 

base, scabrous; blades (5–)17–30 cm long, acuminate or apiculate, sparsely hispid or scabrous below. Foliage leaves: 8–12 per 

branch, pendent; sheath apices glabrous, occasionally with a few 

ciliate hairs, surface glabrous; blades linear to linear-lanceolate, 

3–11.5 cm × 3–6(–12) mm, bases attenuate, margins scabrous, 

apices long-acuminate, glabrous above, scaberulous below; 

pseudopetiolate. Panicles open, pyramidal, 3–8 cm long; rachis 

scaberulous. Spikelets lanceolate, 7–9 mm long.

Evidently restricted to the Andes of Venezuela, where it is 

found in Lara, Mérida, Táchira, and Trujillo; Map 276). In the 

park, common in open areas of cloud forest on both slopes of 

Guaramacal; 2,100–2,900 m.

Chusquea multiramea L. G. Clark & F. Ely is part of a complex of species that includes C. purdieana Munro. As noted by 

Clark and Ely (2013), the former can easily be distinguished from 

the latter when flowering because it has open, pyramidal synflorescences, shorter spikelets, and apically mucronate (never awned) 

lemmas. In vegetative specimens the differences are more subtle, 

C. multiramea differs from C. purdieana by its longer, sparsely 

hispid culm leaf sheaths, which have the margins basally fused for 

a shorter distance from the girdle (0.5–2.5 vs. 3–6.5 cm).

Chusquea multiramea is a dominant element of light gaps 

in cloud forest and in the transition from cloud forest to secondary forest, especially in heavily disturbed areas. In contrast, 

C. purdieana is most often found in upper montane forest and 

MAP 275. Chusquea mollis occurrence in Venezuela. MAP 276. Chusquea multiramea occurrence in Venezuela.
NUMBER 100 • 227

subpáramo. Although C. purdieana occurs in Venezuela (Mérida 

and Zulia states), it is more commonly encountered in the Andes 

of Colombia.

Chusquea serpens L. G. Clark, Ann. Missouri Bot. Gard. 72: 

870, fig. 4. 1985.

Woody bamboos. Culms 10–20 m × 0.5–1 cm, scandent, 

often trailing, stoloniferous; internodes 6–30(–40) cm long, 

solid, green to green mottled with purple, retrorsely scabrous. 

Branching infravaginal. Culm leaves: sheaths ± rectangular, 

6–10 cm long, glabrous or hispid; blades triangular, 10–20 

cm long, bases cordate, margins ciliate, apices setose, glabrous 

above and below, deciduous. Foliage leaves: 3–7 per complement; sheaths keeled, outer margins hairy, glabrous above and 

below; blades ovate to narrowly ovate, 10–30 × 2–5 cm, bases 

rounded-truncate, margins scaberulous, apices tapering to a long 

bristle-like tip, glabrous above and below, sometimes pilose at 

base; pseudopetiolate. Panicles narrow, ± secund, 30–55(–60) cm 

long, rachis glabrous; primary branches appressed, 5–9 cm long 

at base of panicle, progressively shorter toward apex. Spikelets 

lanceolate or ovate, 11–15 mm long, awned.

Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, and Ecuador). In Venezuela, recorded from the Andes (Lara, Mérida, Táchira, and 

Trujillo) and the Cordillera de la Costa (Guárico; Map 277). 

In the park, collected in Qda. Jirajara on the south slope; 

2,500 m.

This species is easily distinguished from the other bamboos 

in the park by its stoloniferous habit and ovate to narrowly ovate 

foliage leaf blades tapering to long bristle-like tips.

Chusquea spectabilis L. G. Clark, in Fisher et al., Syst. Bot. 34: 

681. 2009, nom. nov. Planotia aperta Munro, Trans. Linn. 

Soc. London 26(1): 73. 1868, non Chusquea aperta L. G. 

Clark, Ann. Missouri Bot. Gard. 74: 426, fig. 1E–H. 1987. 

Neurolepis aperta (Munro) Pilg., in Engler and Prantl, Nat. 

Pflanzenfam. Nachtr. 3: 21. 1906; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 57. 2000 [2001].

Woody bamboos. Culms 4–8 m tall, erect, unbranched 

above the base. Sheaths pubescent; auricles erect; ligules membranous, bilobed, scarious, ~4 cm long; blades lanceolate, (0.7–)

2–3 m × 4.5–15 cm, erect, bases attenuate, margins minutely 

strigose, apices attenuate, glabrous above, glaucous below; pseudopetiolate. Panicles ovate, lax, 70–90 cm long, purple; primary 

branches 20–35 cm long; panicle axis and branches glabrous. 

Spikelets oblong, (6–)10–15 mm long.

Found in South America (Colombia, Venezuela, and Ecuador). In Venezuela, known only from the Cordillera de Mérida 

(Mérida and Trujillo; Map 278). In the park, collected in cloud 

forest and subpáramo in Qda. Jirajara on the south slope; 

~2,500–3,000 m.

Among the bamboos found in Guaramacal this species 

is distinguished by the very large size and diffuse aspect of its 

inflorescence. In addition, the species is remarkable in having 

short, obtuse glumes; long, delicate pedicels much longer than 

the spikelets; and an elongated internode between the glumes 

and first sterile lemma. Chusquea spectabilis was long thought 

to be an isolated species, but it shows relationships to two species endemic to the Andes of Colombia: C. petiolata (Davidse & 

L. G. Clark) L. G. Clark and C. silverstonei (Davidse & L. G. 

Clark) L. G. Clark.

MAP 277. Chusquea serpens occurrence in Venezuela. MAP 278. Chusquea spectabilis occurrence in Venezuela.
228 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Chusquea spencei Ernst, J. Bot. 10: 262. 1872; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 599–600, 

fig. 46. 2010.

Shrubby, fastigiate bamboos; forming dense colonies. 

Culms 2–6 m × 0.5–4 cm, erect, tips pendulous; internodes 5–15 

cm long, solid, becoming fistulose, yellow to reddish-brown, 

smooth, waxy. Culm leaves: sheaths ± triangular, 5–20 cm long, 

hispid below; blades narrowly triangular to triangular, 2–6 cm 

long, erect, apices subulate, pubescent above, hispid below. 

Branching intravaginal; central branch robust, all branches erect 

and rebranching vigorously. Foliage leaves: 3–5 per complement; 

sheaths glabrous, margins sometimes ciliate near apex; blades 

linear to linear-lanceolate, 5–15 cm × 2–6 mm, arched or erect, 

bases attenuate, margins slightly cartilaginous, sometimes scabrous, serrated toward the apex, apices setose, glabrous above 

and below; pseudopetiolate. Panicles narrow, congested, 2–12 

cm long; branches appressed, pubescent. Spikelets 4.6–7.3 mm 

long, glabrous.

Found in South America (Colombia and Venezuela). In Venezuela, found in the Andes (Lara, Mérida, Táchira, and Trujillo), 

the Sierra de Perijá (Zulia), and the Cordillera de la Costa (Anzoátegui, Distrito Federal, and Miranda; Map 279). In the park, 

collected in cloud forest and dwarf forest on the south slope and 

in the Páramo de Guaramacal; 2,800–3,100 m.

As noted above, Chusquea spencei and C. angustifolia are 

similar in their foliage leaf blades and spikelets, and discriminating between the two species can be difficult.

Chusquea tessellata Munro, Trans. Linn. Soc. London 26(1): 60. 

1868. Swallenochloa tessellata (Munro) McClure, Smithsonian Contr. Knowl. 9: 113. 1973.

Woody bamboos; forming dense colonies. Culms 2–3 m × 

4–10 mm, erect; internodes 4–8(–10) cm long, solid, becoming fistulose, yellow, smooth, mealy below and sometimes above the leaf 

scar. Culm leaves: sheaths ± rectangular, 5–8.5 cm long, glabrous 

below, persistent; blades triangular, 5–11.5(–18.5) cm long, erect, 

bases slightly cordate, margins ciliate, apices acuminate to shortly 

subulate, glabrous, caducous. Branching intravaginal; branches 

erect and rebranching from base. Foliage leaves: 4–10 per complement; sheaths apical margins pubescent, persistent; ligules internal 

and external, almost always ciliate; blades lanceolate, 5.8–23 × 

0.6–2.6 cm, erect, bases rounded or truncate, margins short-ciliate 

below and ± scaberulous above, apices acuminate-subulate, not 

tessellate (i.e., checkered) above, usually tessellate below, glabrous; 

pseudopetiolate. Panicles contracted, often interrupted near the 

base, 6–30 cm long, dark purple or light brown; primary panicle 

branches appressed, 1–3 cm long; panicle axis and branches pubescent. Spikelets oblong, compressed, 3.9–9 mm long.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). Venezuelan collections are from the Andes 

(Apure, Lara, Táchira, and Trujillo) and the Sierra de Perijá 

(Zulia; Map 280). Our record is based on a sterile collection made 

in shrubby páramo in the Fila de Agua Fría; 2,700–2,800 m.

This species is characterized by culms that are mealy below 

and sometimes also above the leaf scar, and culm leaf sheaths 

that ± equal the slightly cordate sheath blades in length.

Cinna L.

Cinna L., Sp. Pl. 5. 1753.

Perennial plants; caespitose. Culms unbranched, erect. 

Sheaths open, glabrous; ligules eciliate, membranous; blades 

MAP 279. Chusquea spencei occurrence in Venezuela. MAP 280. Chusquea tessellata occurrence in Venezuela.
NUMBER 100 • 229

linear, flat, margins scabrous, scabrous or smooth above and 

below. Inflorescences terminal panicles, spreading or ascending. Spikelets elliptic, laterally compressed, ± obtuse or acute, 

1(2)-flowered, bisexual. Disarticulation below the glumes. 

Rachilla prolonged beyond palea as a minute stub or bristle 

or absent. Glumes equal or subequal, acute, membranous, 

1–3-nerved, sometimes minutely awn-tipped. Lemmas membranous, faintly 3–5-nerved, awned or not (our species); awn short, 

straight. Palea mostly smooth, 1- or 2-nerved. Stamens 1 or 2. 

Styles 2. Caryopses oblong, often beaked; beak formed by the 

persistent style. [Pooideae.]

Four species in temperate regions of North America (Canada and USA), Central America, South America (Colombia, 

Venezuela, Ecuador, Peru, and Bolivia), Europe, and Asia. One 

species occurs in Venezuela.

References. Brandenburg et al. (1991); Gillespie et 

al. (2008, 2010); Tucker (1996).

Brandenburg et al. (1991) define the genus by the following 

combination of morphological characters: 1-flowered spikelets, 

disarticulation below the glumes, awn (when present) subterminal, rachilla typically prolonged behind the palea as a small glabrous or scaberulous stub or bristle, and paleas 1- or 2-nerved 

with the nerves very close together.

In molecular analyses Soreng and Davis (2000) resolved

Cinna as a close relative of Sphenopholis Scribn. and Trisetum

Pers., but that result is now considered to be a laboratory error 

(Gillespie et al., 2008). Quintanar et al. (2007), also using DNA, 

found that Cinna was closely related to a morphologically diverse assemblage of Aveneae that formed a well-supported group 

with Poa L. and relatives. Support for the monophyly of Cinna, 

however, is poor (Gillespie et al., 2010).

Cinna poiformis (Kunth) Scribn. & Merr., Bull. Div. Agrostol. 

U.S.D.A. 24: 21. 1901 (“poæformis”); Briceño, in Morillo 

et al., Bot. Ecol. Monocot. Páramos Venez. 2: 601–602, 

fig. 49. 2010. Deyeuxia poiformis Kunth, in H. B. K., Nov. 

Gen. Sp. [fol.] 1: 119. 1815 [1816]; ibid. [qu.] 1: 146. 1815 

[1816] (“poæformis”).

Culms 60–120(–150) cm tall. Sheaths overlapping, glabrous; ligules prominent, (1–)5–12 mm long; blades 10–20

(–30) × 0.4–0.8 cm, acuminate, glabrous. Inflorescences solitary, 15–45 cm long; branches spreading, pubescent with appressed and scattered hairs. Spikelets borne in dense clusters 

near the branch tips, narrowly ovate, 2–3 mm long, scabrous, 

1-flowered. Glumes equal, lanceolate, 1.8–3.5 mm long, 

3-nerved. Lemmas narrowly ovate, 2–3 mm long, 5-nerved, 

awn usually absent (0.3–0.4 mm long when present). Paleas 

slightly longer than lemmas, 2–3 mm long, 1-nerved. Stamens 

3. Caryopses 1.5–2 mm long.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Ecuador, Peru, and Bolivia) at higher elevations. In Venezuela, found in the Andes (Mérida and Trujillo) 

and the Cordillera de la Costa (Distrito Federal and Miranda; 

Map 281). Páramo del Pumar; ~2,600 m.

Cortaderia Stapf

Cortaderia Stapf, Gard. Chron., ser. 3, 22: 378. [27 Nov] 1897, nom. cons.

Lamprothyrsus Pilg., Bot. Jahrb. Syst. 37, Beibl. 85: 58. 1906.

Perennial plants; gynodioecious, pistillate only or pistillate 

and apomictic; densely caespitose. Culms unbranched, erect, 

clumped. Sheaths open, often overlapping, glabrous or pubescent, often curling and disintegrating with age; ligule a dense 

rim of hairs; blades usually basal, flat or convolute, linear, 

sclerophyllous, margins often scabrid or serrate. Inflorescences 

terminal, plumose panicles, loose or contracted, green, silver, 

brown or pink to purple. Spikelets somewhat compressed laterally, 2–9-flowered, staminate or pistillate (unisexual), sometimes 

bisexual; disarticulation above the glumes and near the base 

of each rachilla segment. Glumes linear-lanceolate, unequal, 

1–5-nerved, apices pointed, rounded or bifid, hyaline or membranous, shorter to longer than florets; lemmas ovate or lanceolate, ± equal to spikelets in length, 3–7-nerved; lemma lobes 

usually absent; setae absent to well developed; central awn not 

differentiated into a twisting column and straight apical part; 

occasionally lemma blade extending to the tip of the awn. Paleas 

lorate to linear, occasionally with tufts of hairs on margins; calli 

rounded to truncate, pilose. Lodicules 2, rhomboid to cuneate, 

ciliate. Anthers (staminate or hermaphroditic flowers) 3. Staminodes (pistillate flowers) 3. Ovary glabrous, rudimentary in 

staminate flowers. Styles (pistillate flowers) 2. Caryopses lorate, 

ellipsoid, turbinate or obovoid. [Danthonioideae.]

A neotropical genus of 20 species found in Central America 

and South America (Colombia, Venezuela, Guyana, Ecuador, 

Peru, Bolivia, Brazil, Chile, Paraguay, Argentina, and Uruguay). 

MAP 281. Cinna poiformis occurrence in Venezuela.
230 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Seven species are found in Venezuela, all restricted to higher 

elevations.

References. Barker et al. (2007); Connor and 

Edgar (1974); Linder et al. (2010); Pirie et al. (2009); Tucker 

(1990).

We follow Linder et al. (2010) in limiting Cortaderia to 

American species and segregating the closely related New Zealand species as Austroderia N. P. Barker & H. P. Linder and the 

sole New Guinea species as Chimaerochloa H. P. Linder. All 

three of these genera share the gynodioecious reproductive system, which is relatively rare in grasses.

Cortaderia selloana (Schult. & Schult. f.) Asch. & Graebn., 

also known as “pampas grass,” is a popular ornamental in North 

America, Europe, Africa, Australia, and New Zealand. Several 

other species of Cortaderia also are cultivated, and a number of 

these have become invasive weeds.

Cortaderia hapalotricha (Pilg.) Conert, Syst. Anat. Arundineae 

102. 1961; Briceño, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 2: 603, fig. 50, foto 8. 2010. Danthonia 

hapalotricha Pilg., Bot. Jahrb. Syst. 25: 715. 1898.

FIGURE 26C

Cortaderia roraimensis auct., non N. E. Br. (Pilg.); Dorr et al., Contr. U.S. 

Natl. Herb. 40: 57. 2000 [2001].

Plants pistillate and apomictic. Culms to 1.5 m tall. Basal 

leaves numerous, stiffly erect; sheaths longer than internodes, 

purple-tinged above; blades linear, 20–40(–90) cm × 3–5(–10) 

mm, convolute, apices tapering to points, margins scabrous, 

densely silky above (above the ligule), glabrous below. Panicles 

(10–)20–35 cm long, pink to purple. Spikelets pistillate or perfect, 

with 2–5 florets. Glumes narrowly lanceolate, subequal to equal, 

~12–18 mm long, 1-nerved, slightly bifid; lemmas 5–8 mm long 

(excluding awns), lanceolate with lateral teeth and a central awn, 

awn geniculate or loosely twisted, 7–15 mm long, 5–7-nerved; 

palea 4–6 mm long, ciliate. Anthers aborted (apomictic flowers) 

or reduced and sterile (pistillate flowers).

Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, Ecuador, Peru, and Bolivia). In Venezuela, found in the Andes (Apure, Barinas, Lara, 

Mérida, Táchira, and Trujillo), the Sierra de Perijá (Zulia), and 

the Cordillera de la Costa (Distrito Federal and Miranda; Map 

282). In the park, this species forms dense colonies in open areas 

of subpáramo and páramo; 2,800–3,150 m.

Cortaderia columbiana (Pilg.) Pilg. could be confused with 

C. hapalotricha. The two species are sometimes sympatric, but 

the former tends to be found at lower elevations in the Venezuelan Andes, whereas the latter is more common in páramo above 

3,000 m elevation. The panicle of C. columbiana is brown, 

whereas that of C. hapalotricha is pink to purple. Also, hermaphroditic florets of C. columbiana are fully fertile, whereas 

those of C. hapalotricha are apomictic with large whitish anthers 

that are empty (i.e., infertile).

Danthonia DC.

Danthonia DC., in Lamarck and A. P. de Candolle, Fl. Franç., 3rd ed., 3: 32. 

1805, nom. cons.

Perennial plants; caespitose, sometimes shortly rhizomatous; culms simple, erect; often with cleistogenes (i.e., reduced 

cleistogamic inflorescences consisting of solitary, sessile, single 

flowers with lemma and palea but without outer empty glumes) 

in the axils of upper leaves. Sheaths open to the base; auricles absent; ligules ciliate; blades flat or involute, glabrous or pubescent. 

Inflorescences terminal panicles, racemes or a solitary spikelet; 

spikelets terete or laterally compressed, with 2–12 florets, the 

terminal floret reduced; disarticulation usually below the florets. 

Glumes subequal or the lower glumes a little longer than the upper 

ones, shorter to longer than the florets, chartaceous, 3–9-nerved; 

calli rounded or truncate, villous; lemmas 5–9-nerved, apically 

3-toothed to lobed; lemma lobes acute, shorter to longer than 

the lemma body, usually prolonged into a terminal seta; lemma 

central awn exceeding the lobes, geniculate and twisted below 

the geniculation, straight and hairlike toward the apex; paleas 

ligulate to linear; lodicules 2, bristly; anthers 3; ovary glabrous. 

Caryopses ligulate, ellipsoid to obovoid. [Danthonioideae.]

A genus of 25 species, 23 found in temperate parts of the 

Americas and 2 in Europe. In the Americas, Danthonia occurs in 

North America (Canada and USA), Central America, the Greater 

Antilles, and South America (Colombia, Venezuela, Ecuador, 

Peru, Bolivia, Brazil, Chile, Argentina, and Uruguay). A single 

species is known from páramo and subpáramo in Venezuela.

References. Baeza P. (1996); Barker et al. (2007); 

Dertscheny (1995); Linder et al. (2010); Pirie et al. (2009); 

Tucker (1990).

Although the Danthonioideae form a well-defined clade of 

mostly Southern Hemisphere grasses, generic limits within the 

MAP 282. Cortaderia hapalotricha occurrence in Venezuela.
NUMBER 100 • 231

subfamily have been unstable (Linder et al., 2010). We have adopted here a narrower concept of Danthonia than has generally 

been used in the Americas and we exclude species from Africa, 

Australasia, and Malesia, most of which now are placed in related genera (Linder et al., 2010).

Danthonia secundiflora J. Presl, Reliq. Haenk. 1: 255. 1830; 

Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 2: 607, fig. 54. 2010.

The nominate subspecies is the only one found in Guaramacal and in Venezuela.

Danthonia secundiflora subsp. secundiflora

Culms 20–45(–70) cm tall, glabrous. Sheaths glabrous except 

for oral hairs; blades filiform, 5–25(–55) × 0.1–0.4 cm, involute, 

glabrous or occasionally with short hairs above. Inflorescences 

panicles, 5–15 cm long, ± lax. Spikelets composed of 3–7(–9) 

fertile florets, elliptic, 7–20 mm long, laterally compressed. Calli 

evident, bearded. Glumes subequal, 10–15(–20) mm long, equal 

to or longer than the florets, membranous, 3-nerved. Lemmas 

elliptic, 1.5–3 mm long, chartaceous, 5-nerved, 3-awned, central 

awn arising from a sinus, geniculate, 10–12 mm long, twisted, 

lateral awns 3–4 mm long. Paleas oblanceolate. Lodicules 2, cuneate, fleshy, glabrous. Ovary ~1 mm long. Caryopses ellipsoid, 

1.5–2.5 mm long.

Found in Mexico, Central America, and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Argentina, and 

Uruguay). In Venezuela, recorded from higher elevations in the 

Andes (Apure, Lara, Mérida, Táchira, and Trujillo), the Sierra 

de Perijá (Zulia), and the Cordillera de la Costa (Distrito Federal 

and Miranda; Map 283). Páramo de Vicuyal; 2,700–2,800 m.

In addition to the nominate subspecies, which has an extensive 

range, Baeza P. (1996) recognized Danthonia secundiflora subsp. 

charruana (Swallen) Roseng. et al. and D. secundiflora subsp. matthei C. M. Baeza, these latter two subspecies occurring only in 

southern South America (Bolivia, Brazil, and Paraguay). The three 

subspecies are distinguished by relatively minor characters of the 

lemmas (length of awns and pubescence) and calli (length of hairs).

Dichanthelium (Hitchc. & Chase) Gould

Dichanthelium (Hitchc. & Chase) Gould, Brittonia 26: 59. 1974. Panicum

subgen. Dichanthelium Hitchc. & Chase, Contr. U.S. Natl. Herb. 15: 

20, 142. 1910.

Perennial plants; caespitose, with or without a basal rosette of 

shorter, broader leaves than those found on culms. Culms erect, ascending or decumbent, herbaceous, simple or branched; internodes 

hollow. Cauline leaves usually longer and narrower than basal 

leaves; sheaths pilose or glabrous, conspicuous glands present or 

not; ligules a ring of hairs, membranous, membranous and ciliate, 

or sometimes absent; blades linear to ovate-lanceolate, densely pilose to glabrous, flat or involute. Inflorescences terminal and axillary panicles, lax or occasionally contracted; disarticulation below 

the glumes. Spikelets ellipsoid to obovoid, awnless; cleistogamous 

spikelets present or not. Glumes unequal, (1–)7–15-nerved, membranous; upper glumes and lower lemmas similar, herbaceous, 

nearly as long as spikelets or lower lemmas slightly shorter than 

spikelets; lower florets usually sterile, rarely staminate; upper florets bisexual, coriaceous, glossy, glabrous; lower lemmas similar 

to upper glumes; lower paleas shorter than or nearly equal to the 

lemmas; lodicules 2; anthers 3; styles 2. [Panicoideae.]

An American genus of 50–75 species found in North America (Canada and USA), Mexico, the West Indies, and South 

America (all country-level political units). Thirteen species are 

recorded from Venezuela.

References. Aliscioni et al. (2003); Freckmann 

(1981); Gould and Clark (1978); Hammer (2010); Morrone et 

al. (2012); Zuloaga (1987); Zuloaga et al. (1993).

Dichanthelium, originally circumscribed as a subgenus of 

Panicum L., was created to accommodate a group of species with 

a distinctive leaf dimorphism and breeding system. As noted by 

Zuloaga et al. (1993), plants of this genus overwinter with a 

rosette of short, broad leaves that produce simple culms with 

terminal, chasmogamous inflorescences in the spring. Later in 

the season culms bear fascicles of reduced leaves and branches 

together with cleistogamous inflorescences.

Dichanthelium acuminatum (Sw.) Gould & C. A. Clark, Ann. 

Missouri Bot. Gard. 65: 1121. 1978 [1979]. Panicum acuminatum Sw., Prodr. 23. 1788.

Guaramacal material can be referred to the nominate variety.

Dichanthelium acuminatum var. acuminatum

Culms usually in large clumps, 10–45(–90) cm tall, branching from the middle and upper nodes; nodes glabrous to pilose; 

MAP 283. Danthonia secundiflora subsp. secundiflora occurrence 

in Venezuela.
232 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

internodes (2–)4–7(–9) cm long, glabrous to pilose. Sheaths 

usually shorter than internodes, glabrous or densely pubescent, margins ciliate; ligules ciliate, hairs 2–6 mm long; blades 

linear-lanceolate, lanceolate to ovate, 4–11 × 0.3–1 cm, flat, 

bases rounded to subcordate, long-hispid (hairs ≤1.5 mm long) 

to glabrous above, pilose below. Inflorescences terminal panicles, 3–9 × 3–9 cm, lax; main axis flexuous, glabrous to pilose. 

Spikelets 1.5–2.5 mm long, apically subacute, puberulent; lower 

glumes ovate-acuminate, 0.5–1.1 mm long; upper glumes ovate, 

~1.5 mm long, 9-nerved; lower lemmas resembling glumes, 

7–9-nerved; lower paleas small; upper anthecium (i.e., fruit derived from a spikelet with 1 floret) ovoid.

Found in North America (Canada and USA), Mexico, Central America, the West Indies, and South America (Colombia, 

Venezuela, and Ecuador); introduced in Asia and Hawaii. In 

Venezuela, found in the Cordillera de Mérida (Mérida and Trujillo) and the Cordillera de la Costa (Aragua, Distrito Federal, 

and Miranda; Map 284). Collected in El Campamento in Qda. 

Jirajara on the south slope of Guaramacal; 2,000 m.

Dichanthelium acuminatum is a polymorphic and taxonomically difficult species complex (Gould and Clark, 1978; Freckmann, 

1981; Zuloaga et al., 1993; Hammer, 2010). Our material keys to 

the nominate variety in the revision by Zuloaga et al. (1993).

Didymogonyx (L. G. Clark & Londoño) 

C. D. Tyrrell et al.

Didymogonyx (L. G. Clark & Londoño) C. D. Tyrrell et al., in Tyrrell et al., 

Molec. Phylogenet. Evol. 65: 146. 2012. Rhipidocladum sect. Didymogonyx L. G. Clark & Londoño, Amer. J. Bot. 78: 1271. 1991.

Large, woody bamboos; caespitose. Culms erect at base, 

arched toward apex; primary element of the branch complement 

at midculm nodes flat, unsegmented, appressed and adnate to 

the culm surface; second-order branches flabellate; internodes 

smooth, hollow, each long internode alternating along the length 

of the culm with 1 short internode. Culm leaves: coriaceous, well 

developed or not, glabrous, ciliate or pubescent: sheaths glabrous 

or hispid and glabrescent, bearing setose auricles or fimbriae at 

apices, as long as or longer than the blade; inner ligules truncate, 

ciliate or conspicuously fimbriate along the margin; blades triangular, pubescent between the nerves above, hispid-pubescent and 

glabrescent below, junction with the sheath almost symmetrical, 

± horizontal, slightly curved at the middle. Branching intravaginal, restricted to middle and upper portions of culms; secondary branches slender, numerous, 20–30(–50) per node. Foliage 

leaves: sheaths glabrous, not auriculate, apically fimbriate; fimbriae free, basally straight, distally wavy to curly; blades linear, 

7–17(–19) × 0.3–1.2(–2.1) cm, bases rounded-attenuate, margins serrulate or scabrous, apices attenuate-acuminate, glabrous 

or scabrid above, glabrous below. Inflorescences racemiform or 

subpaniculiform, 10–25(–40) cm long, with 10–20(–30) spikelets 

per synflorescence. Spikelets slightly compressed, composed of 

4–9(–12) florets and 1 terminal rudimentary floret, 3(–5) glumes, 

and 1 sterile lemma; glumes triangular to narrowly ovate, shortly 

mucronate at the tip; fertile lemmas ovate or narrowly ovate, 

shortly mucronate, glabrous or scabrous; paleas 2-keeled, keels 

ciliolate; lodicules 3; anthers 3; stigmas 2. [Bambusoideae.]

A neotropical genus of 2 species found in the Andes of 

South America (Venezuela and Colombia). One species is present in Venezuela.

References. Clark and Londoño (1991); Tyrrell et 

al. (2012).

Molecular data (Tyrrell et al., 2012) support the monophyly 

of this small genus of arthrostylidioid bamboos, which is related 

to Arthrostylidium and Rhipidocladum. The two species of Didymogonyx were included most recently in Rhipidocladum s.l., 

albeit in a section readily distinguished from Rhipidocladum s. 

str. by having alternating long and short internodes along the 

culm, culm leaf fimbriae, spikelets in pairs and triads, and several 

other distinctive morphological and foliar anatomical characters 

(Clark and Londoño, 1991; Tyrrell et al., 2012).

Didymogonyx geminatum (McClure) C. D. Tyrrell et al., in 

Tyrrell et al., Molec. Phylogenet. Evol. 65: 146. 2012. Arthrostylidium geminatum McClure, J. Wash. Acad. Sci. 32: 

169, fig. 2. 1942. Rhipidocladum geminatum (McClure) 

McClure, Smithsonian Contr. Bot. 9: 105. 1973; Dorr et 

al., Contr. U.S. Natl. Herb. 40: 58. 2000 [2001]; Briceño, in 

Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 678, 

fig. 108. 2010.

Culms 6–10(–15) m tall, 1–5 cm in diameter; long internodes 0.5–1 m long, short internodes 5–10 mm long. Culm 

leaves: sheaths (9–)11–20 × 4–9 cm, glabrous or hispid and 

glabrescent, apices fimbriate, margins ciliate or smooth; inner 

MAP 284. Dichanthelium acuminatum var. acuminatum occurrence in Venezuela.
NUMBER 100 • 233

ligules 0.5–2 mm long, truncate, glabrous, marginally ciliate; 

blades triangular, (3–)5–12 ×1.5–6 cm, margins glabrous to 

strigose, pubescent between the nerves above, hispid-pubescent 

and glabrescent below. Foliage leaves: 3–7(–8) per branch; 

outer ligules 0.2–0.4 mm long, truncate, glabrous, margins ciliolate; inner ligules 0.1–0.4 mm long, truncate; blades linear to 

linear-lanceolate, 5–11(–17) × 0.3–1.8(–2.1) cm, bases roundedattenuate, ± oblique, margins minutely serrulate, apices acuminate, glabrous above and below; pseudopetioles 1–4(–6) mm 

long, twisted. Inflorescences subpaniculiform, 8–15 cm long, 

with 15–25(–30) spikelets. Spikelets 0.4–1(–2.7) cm long, with 

4(–5) florets and 1 terminal rudimentary floret; disarticulation 

between fertile florets. Glumes 3(–5); lower glumes 2–4 mm 

long, 1–3-nerved; upper glumes 3–5 mm long, 5–7-nerved; sterile lemmas 4–7 mm long, 7-nerved; fertile lemmas 6–10 × 3–4 

mm, 7-nerved; paleas 5–7 mm long.

Endemic to South America (Colombia and Venezuela). In 

Venezuela, restricted to the Andes (Lara, Mérida, Táchira, and 

Trujillo; Map 285). In the park, found in forest on both slopes of 

Guaramacal; (1,500–)2,400–2,600 m.

This species was described from material collected in the 

nearby Páramo de la Cristalina (Trujillo state).

A report of this species occurring in Ecuador is based on 

misidentification of material of Aulonemia hirtula (Pilg.) McClure (Judziewicz in Jørgensen and León-Yánez, 1999).

Eragrostis Wolf

Eragrostis Wolf, Gen. Pl. 23. 1776.

Annual or perennial plants; caespitose, rhizomatous or stoloniferous. Culms erect, decumbent or geniculate, sometimes 

rooting at lower nodes; internodes solid or hollow. Sheaths open, 

apices with tufts of hairs; ligules membranous and ciliate or ciliate; blades filiform to linear, flat, folded or involute, narrow. Inflorescences terminal panicles, open to contracted or spikelike. 

Spikelets pedicellate, almost always laterally compressed, with 

2–60 florets; disarticulation below fertile florets or sometimes 

below glumes. Glumes ± equal in size and shape, membranous 

or papery, hyaline, 1(–3)-nerved, usually glabrous and keeled; 

lemmas similar or very similar to the glumes, 3-nerved; paleas 

shorter than lemmas, almost always hyaline, 2-keeled, keels 

winged or ciliate; lodicules 2(0); anthers 2 or 3. Caryopses lenticular or subellipsoid. [Chloridoideae.]

A cosmopolitan genus of ~350 species with its greatest diversity in subtropical regions (low elevations) and in the tropical 

Andes (high elevations). Approximately 75 species occur in the 

Americas, including North America (Canada and USA), Central 

America, the West Indies, and South America (all upper-level political units). In Venezuela, ~29 species have been reported.

References. Giraldo-Cañas et al. (2012); Graterol 

et al. (1989); Peterson and Giraldo-Caños (2008); Peterson et 

al. (1997).

• Eragrostis tenuifolia (A. Rich.) Hochst. ex Steud., Syn. Pl. Glumac. 1: 268. 1854. Poa tenuifolia A. Rich., Tent. Fl. Abyss. 

2: 425. 1850.

Eragrostis tenuifolia Hochst., in Schimper, Flora 24(Intell.1): 20. 1841, 

nom. nud.

Culms 20–50(–75) cm tall, unbranched or branched from 

basal nodes. Sheaths strongly compressed and keeled, glabrous, 

apices pilose; ligules ciliate; blades flat or folded, 5–25 cm × 

1–3 mm, glabrous to scaberulous above, glabrous below. 

MAP 286. Eragrostis tenuifolia occurrence in Venezuela.

MAP 285. Didymogonyx geminatum occurrence in Venezuela.
234 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Inflorescences solitary panicles, 15–25 × 3–10 cm, pyramidal; 

pulvini of the primary branches densely ciliate. Spikelets linear, 

3–6 mm long, with 6–14 florets. Florets 3–10 mm long, pedicels 

slightly scabrous; glumes reduced, not keeled, equal or subequal, 

to 1 mm long; lemmas ovate to acute; paleas smaller than lemmas. Caryopses ovoid, strongly flattened laterally, ~1 mm long.

Native to Africa and Asia, this species expanded its range 

rapidly after being introduced into the Americas, where it is now 

found in Mexico, Central America, the West Indies, and South 

America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, 

Chile, and Argentina). In Venezuela, presently known only from 

the Andes (Mérida, Táchira, and Trujillo; Map 286). In the park, 

found near the Laguna de Aguas Negras; 1,900 m.

Eriochrysis P. Beauv.

Eriochrysis P. Beauv., Ess. Agrostogr. 8. 1812.

Perennial plants; caespitose. Culms erect. Sheaths cylindrical, pubescent; ligules membranous, ciliate; blades flat and linear 

to linear-lanceolate, usually densely pubescent. Inflorescences terminal, cylindrical panicles with numerous, short, erect branches, 

usually densely villous-silky, yellowish to brown. Spikelets 

paired at each node of the rachis, subequal, dorsally compressed, 

lanceolate, awnless; 1 sessile and 1 short-pedicellate; sessile 

spikelets bisexual; disarticulation at the base of the sessile spikelets, the 2 spikelets of a pair falling together or the pedicellate 

one dropping from the pedicel; pedicellate spikelets pistillate; 

glumes equal, chartaceous, concealing the florets; lower glume 

flat dorsally, margins inflexed and covering upper glume; upper 

lemma membranous or hyaline, entire, muticous or mucronate; 

sterile lemmas and fertile lemmas nerveless, slightly shorter than 

the glumes; palea absent. Lower floret sterile, upper floret bisexual in sessile spikelets and pistillate in pedicellate spikelets. 

[Panicoideae.]

A cosmopolitan tropical and subtropical genus of 10 species. In the Americas, found in Mexico, Central America, the 

Greater Antilles, and South America (Colombia, Venezuela, the 

Guianas, Peru, Bolivia, Brazil, Paraguay, Argentina, and Uruguay). Two species occur in Venezuela.

Reference. Swallen (1966).

Eriochrysis cayennensis P. Beauv., Ess. Agrostogr. 8, t. 4, fig. 11. 

1812 (“Cayanensis”); Judziewicz, in Berry et al., Fl. Venez. 

Guayana 8: 106. 2004.

Culms 0.5–1.5(–3) m tall. Basal leaves numerous; sheaths 

glabrous except auricles densely pubescent; ligules membranous; 

blades 1.5–4.5 × 0.1–0.8(–1.2) cm, densely villous above and 

below. Inflorescences (including peduncles) to 30–50 × 1.2–2.5 

cm, soft to the touch, chestnut or coppery-brown. Sessile spikelets lanceolate, 2–3 mm long; pedicellate spikelets 1.5–2 mm 

long. Caryopses obpyriform, ~1 mm long.

Found in Mexico, Central America, the Greater Antilles, 

and South America (Colombia, Venezuela, Trinidad and Tobago, the Guianas, Peru, Bolivia, Brazil, Paraguay, Argentina, 

and Uruguay). Widely distributed in Venezuela (Amazonas, Anzoátegui, Apure, Aragua, Bolívar, Guárico, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, and Trujillo; Map 287). In 

the park, found in open, disturbed sites including the track that 

leads to El Campamento and Qda. Honda; 1,800–2,000 m.

This distinctive grass is recognized easily by its long chestnut or coppery-brown inflorescences that superficially resemble 

small “cat tails” of the genus Typha L. (Typhaceae).

Festuca L.

Festuca L., Sp. Pl. 73. 1753.

Perennials; caespitose, rhizomatous or occasionally stoloniferous. Culms unbranched, usually glabrous, sometimes scabrous 

below the inflorescences. Leaves cauline or basal; sheaths with 

margins free or partially united, sometimes auriculate; ligules 

membranous, sometimes coriaceous, generally ciliate; blades 

flat or conduplicate to involute, linear, apices acuminate, never 

navicular. Inflorescences panicles, contracted or lax, sometimes 

reduced to racemes. Spikelets compressed laterally, 2- to manyflowered, florets bisexual, pedicellate; rachillae glabrous; disarticulation above the glumes and between the florets. Glumes 2, 

subequal, usually shorter than adjacent lemmas; lower glumes 

1(3)-nerved, upper glumes 3(5)-nerved, entire, strongly mucronate or awned; lemmas lanceolate or acuminate, keeled to 

rounded dorsally, 3–7-nerved, usually awned; awn apical or 

subapical; paleas usually shorter than lemmas, membranous, 

scabrous or ciliate, awnless; lodicules 2, often bilobed apically; 

anthers 3; ovary glabrous or hispidulous apically; styles 2, apical. Caryopses obovoid-oblong, usually free of the lemmas and 

paleas. [Pooideae.]

MAP 287. Eriochrysis cayennensis occurrence in Venezuela.
NUMBER 100 • 235

A genus of 450–500 species found in boreal and temperate 

regions of America, Europe, Africa, Madagascar, Asia, Australia, 

and the Pacific Islands. In the Americas, found in North America 

(Greenland, Canada, and USA), Central America, and South 

America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, 

Chile, Paraguay, Argentina, and Uruguay). Eleven species occur 

in Venezuela, all above 2,000 m elevation.

References. Catalán et al. (2004); Stančík (2003, 

2004); Stančík and Peterson (2007); Torrecilla and Catalán 

(2002); Tucker (1996).

The traditional concept of Festuca is likely to change because molecular data indicate that Lolium L. and Vulpia C. C. 

Gmel. are nested within Festuca s.l. (Catalán et al., 2004). Within 

this large clade two separate lineages have been recognized (Torrecilla and Catalán, 2002), which are informally named the 

“broad-leaved” and “fine-leaved” fescues. The former lineage, 

which includes Lolium, has weak support, whereas the latter, 

which includes Vulpia, is well supported.

♦ Festuca guaramacalana Stančík, Novon 14: 343, fig. 1a–e. 

2004; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 620–623, fig. 65. 2010.

Rhizomatous plants, forming small tussocks. Culms 

60–90(–130) cm tall, erect, glabrous. Sheaths membranous to 

coriaceous, striate; auricles absent; ligules 1–1.5 mm long, membranous, sometimes coriaceous, apices acute; blades conduplicate 

to involute, 30–40 × 0.6–0.7 cm, glabrous below. Inflorescences 

panicles, 15–20 × 2–5 cm, flexuous, pendant. Spikelets narrowly 

lanceolate, 8–15 mm long, with 5–7 florets; rachillae pilose; 

glumes 3–7 mm long, narrowly lanceolate, membranous to coriaceous, sparsely scabrous, apices acute; lower glumes 3–4(–5) 

mm long, 1-nerved; upper glumes 7–8 mm long, 3-nerved; lemmas 7–10 mm long, membranous to coriaceous, 5-nerved, apices 

entire, mucronate or shortly awned; calli sparsely pilose; paleas 

finely pilose, almost as long as the lemmas; lodicules ovate, 

2-dentate.

Endemic to Venezuela, where it is found only in the Cordillera de Mérida (Trujillo; Map 288). Evidently endemic to the 

park, where it has been found in the Páramo de Guaramacal and 

the Páramo del Pumar; 2,600–2,900 m.

Festuca dinirica Stančík, endemic to Dinira National Park 

in Lara to the north of Guaramacal, can be distinguished from 

our species by its habit. The former species is caespitose (vs. rhizomatous), and it forms large (vs. small) tussocks. Festuca elviae

B. Briceño, which is found in Mérida state to the south and west 

of Guaramacal, can be distinguished from F. guaramacalana by 

its shorter lower glumes that are 0.8–2.5 mm (vs. >2.5 mm) long.

Stančík (2004) placed Festuca guaramacalana in F. sect. Subulatae Tzvel., a section that corresponds to the broad-leaved 

fescues, but this morphological classification might not coincide 

with a molecular one. Catalán et al. (2004) found that three 

Andean taxa that they sampled, including F. elviae (also placed 

in sect. Subulatae on the basis of morphology), were related 

within the fine-leaved fescue clade, and they concluded that sect. 

Subulatae represented one of several polyphyletic groups of the 

broad-leaved fescues.

Homolepis Chase

Homolepis Chase, Proc. Biol. Soc. Wash. 24: 146. 1911.

Annual or perennial plants; stoloniferous or decumbent, 

rarely caespitose. Culms erect, ascending or decumbent; rooting 

from lower nodes. Sheaths carinate, glabrous or slightly puberulent; ligules membranous, ciliate; blades linear or lanceolate. Inflorescences terminal panicles, open, lax to contracted; spikelets 

broadly ellipsoid, obovoid or narrowly lanceoloid, compressed, 

with 2 florets, disarticulating entire; glumes subequal, ± equal in 

length to spikelet, usually glabrous, 5–9-nerved; lower floret sterile or staminate, lemma glume-like, several-nerved, viscid or not, 

palea well developed; upper floret bisexual, ellipsoid, shorter than 

spikelet, glabrous, lemma often bearded at apex. [Panicoideae.]

A neotropical genus of 5 species found in Mexico, Central 

America, the West Indies, and South America (Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina, and Uruguay). Three species occur in Venezuela.

References. Morrone et al. (2012); Webster et al. 

(1989); Zuloaga and Soderstrom (1985).

Homolepis glutinosa (Sw.) Zuloaga & Soderstr., Smithsonian 

Contr. Bot. 59: 19. 1985; Judziewicz, in Berry et al., Fl. 

Venez. Guayana 8: 118, fig. 80. 2004. Panicum glutinosum

Sw., Prodr. 24. 1788; Dorr et al., Contr. U.S. Natl. Herb. 40: 

57. 2000 [2001].

Perennial plants. Culms decumbent, 0.5–1.5 m tall; internodes 5–20 cm long, hollow. Sheaths shorter than internodes, 

MAP 288. Festuca guaramacalana occurrence in Venezuela.
236 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

papillose-pilose to glabrescent; ligules membranous with short 

hairs; blades oblong-linear to lanceolate, 10–35 × 0.5–2 cm, flat, 

apices attenuate, margins scaberulous below, bases narrowed, 

papillose-pilose to glabrescent above and below, glaucous. Inflorescences lax, diffuse, 15–25 × 10–25 cm; branches spreading, 

lower branches whorled, upper branches alternate and opposite, 

axils of branches densely pubescent. Spikelets obovoid to ellipsoid, 2–3 mm long, viscid at maturity; lower glume 5–9-nerved, 

glabrous; upper glume, 7–9-nerved, glabrous; lower lemma 

glume-like, 5–7-nerved, scabrous; lower palea smaller than lemmas, glabrous; staminate flower absent. Upper anthecium ellipsoid, ~2.5 mm long, apex uncinate.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Guyana, Ecuador, Peru, 

Bolivia, Brazil, Paraguay, Argentina, and Uruguay). In Venezuela, 

reported from Anzoátegui, Aragua, Bolívar, Carabobo, Cojedes, 

Distrito Federal, Falcón, Guárico, Lara, Miranda, Monagas, Portuguesa, Sucre, Táchira, Trujillo, and Yaracuy (Map 289). In the 

park, found along trails and road cuts in montane forest near the 

Laguna de Aguas Negras and in Qda. Segovia; 2,000–2,350 m.

Ichnanthus P. Beauv.

Ichnanthus P. Beauv., Ess. Agrostogr. 56. 1812.

Annual or perennial plants; caespitose, rhizomatous or 

stoloniferous. Culms erect to decumbent, simple or branched, 

glabrous or pubescent. Sheaths glabrous or pilose; ligules membranous; blades flat, linear to ovate, usually lanceolate and asymmetrically narrowed at the base. Inflorescences terminal or axillary, 

simple to compound, panicles; branches diverging, ascending. 

Spikelets usually lanceoloid, laterally compressed, glabrous or 

pubescent, with 2 florets; glumes prominently keeled, 3–9-nerved; 

lower glumes >1⁄2 the length of spikelets; upper glumes and lower 

lemmas ± equal and exceeding in length the upper fertile florets; 

lemmas enclosing paleas and often staminate flowers; fertile florets hardened; margins of upper lemmas flat, rachilla subtending 

lemma minutely stipitate and bearing on either side membranous, 

winglike appendages adnate to the base of the lemma and free 

above, sometimes appendages reduced to swellings; lodicules 2. 

Caryopses ovoid to ellipsoid. [Panicoideae.]

A mostly neotropical genus of ~30 species; 1 species is pantropical. In the Americas, found in Mexico, Central America, the 

West Indies, and South America (Colombia, Venezuela, Trinidad 

and Tobago, the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, 

Argentina, and Uruguay). Sixteen species occur in Venezuela.

References. Boechat (2005); Morrone et al. (2012); 

Stieber (1982, 1987); Webster et al. (1989).

KEY TO THE SPECIES OF ICHNANTHUS

1a. Leaf blades ovate to ovate-lanceolate, 1–3 cm wide; panicles scarcely exserted from the leaf sheaths . . . . . . . . I. pallens

1b. Leaf blades lanceolate or linear-lanceolate, 0.3–0.9(–1.5) cm wide; panicles well exserted from the leaf sheaths . . . . . 2

2a. Spikelets narrowly ellipsoid or lanceoloid, sparsely to densely pilose especially on margins of glumes . . . . I. tenuis

2b. Spikelets ovoid to ovoid-acuminate, glabrous to sparingly pilose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. nemorosus

Ichnanthus nemorosus (Sw.) Döll, in Martius, Fl. Bras. 2(2): 

289. 1877; Judziewicz, in Berry et al., Fl. Venez. Guayana 

8: 125. 2004. Panicum nemorosum Sw., Prodr. 22. 1788.

Annual or perennial plants. Culms creeping, branched, 

rooting at the nodes, nodes often pilose; flowering shoots 30–40 

cm tall. Leaf blades linear-lanceolate, 3.5–6(–8.5) × 0.4–0.6 cm, 

bases asymmetrically subcordate, glabrous to pilose above and 

below. Inflorescences terminal and axillary, well exserted from 

the leaf sheaths, 2–5(–10) cm long, sparsely branched. Spikelets 

2–4 mm long, plump, ovoid to ovoid-acuminate, glabrous to 

sparingly pilose.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Trinidad and Tobago, 

French Guiana, Ecuador, Peru, Bolivia, Paraguay, and Argentina). Widespread in Venezuela (Amazonas, Aragua, Carabobo, 

Cojedes, Distrito Federal, Falcón, Lara, Mérida, Miranda, Portuguesa, Táchira, Trujillo, Yaracuy, and Zulia; Map 290). In the 

park, found on road cuts and in understory vegetation on both 

slopes of Guaramacal; 1,800–2,100 m.

Boechat (2005) excluded this species from Brazil while noting that it was collected on Cerro de la Neblina, which straddles 

the Venezuelan–Brazilian frontier.

MAP 289. Homolepis glutinosa occurrence in Venezuela.
NUMBER 100 • 237

Ichnanthus pallens (Sw.) Munro ex Benth., Fl. Hongk. 414. 

1861; Judziewicz, in Berry et al., Fl. Venez. Guayana 8: 125. 

2004. Panicum pallens Sw., Prodr. 23. 1788.

Panicum pallens var. majus Nees, Fl. Bras. Enum. Pl. 2(1): 137. 1829. Ichnanthus pallens var. majus (Nees) Stieber, Syst. Bot. 12: 207. 1987.

Perennial plants. Culms spreading to creeping, to 1(–3) 

m long, branched, rooting at the nodes, nodes glabrous to pubescent; flowering shoots 15–20 cm tall. Leaf blades ovate to 

ovate-lanceolate, 5–10 × 1–3 cm, bases asymmetrically cordateclasping, sparsely pilose above and below. Inflorescences terminal and axillary, scarcely exserted from the leaf sheaths, 5–10 cm 

long, sparsely branched. Spikelets 3–6 mm long, ± plump, ovoid 

to lanceoloid, glabrous to short pubescent.

Found in Mexico, Central America, the West Indies, South 

America (Colombia, Venezuela, Curaçao, Trinidad and Tobago, 

the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina, 

and Uruguay), tropical Africa, Asia, and Australia. Widespread 

in Venezuela (Amazonas, Anzoátegui, Apure, Aragua, Barinas, 

Bolívar, Cojedes, Delta Amacuro, Distrito Federal, Falcón, Lara, 

Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, 

Trujillo, Yaracuy, and Zulia; Map 291). In the park, found in forest understory and in open areas on the north slope of Guaramacal; 1,900 m.

Stieber (1987) recognized two varieties of Ichnanthus pallens mostly on the basis of spikelet size. Our material could be 

referred to the nominate variety. However, given that the varieties he recognized do not segregate geographically and that he 

noted morphological intermediates, we prefer to recognize a 

single polymorphic species with a cosmopolitan distribution.

Ichnanthus tenuis (J. Presl) Hitchc. & Chase, Contr. U.S. Natl. 

Herb. 18: 334. 1917; Judziewicz, in Berry et al., Fl. Venez. 

Guayana 8: 131, fig. 88. 2004. Oplismenus tenuis J. Presl, 

in C. Presl, Reliq. Haenk. 1: 319. 1830.

Annual plants. Culms spreading to creeping, 10–40 cm long, 

freely branched, branches erect or ascending, rooting at lower 

nodes, nodes glabrous or slightly pubescent. Leaf blades lanceolate, 1–7 × 0.3–0.9(–1.5) cm, bases asymmetrically rounded or 

subcordate, pubescent or pilose above and below. Inflorescences 

MAP 290. Ichnanthus nemorosus occurrence in Venezuela.

MAP 291. Ichnanthus pallens occurrence in Venezuela. MAP 292. Ichnanthus tenuis occurrence in Venezuela.
238 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

terminal and axillary, 2–7(–15) cm long, well exserted from the 

leaf sheaths; branches spreading or ascending. Spikelets 3–5 mm 

long, narrowly lanceoloid.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Trinidad and Tobago, the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, and Argentina); exotic in the 

Greater Antilles. Widely distributed in Venezuela (Amazonas, 

Anzoátegui, Aragua, Bolívar, Cojedes, Distrito Federal, Falcón, 

Lara, Mérida, Miranda, Portuguesa, Sucre, Táchira, Trujillo, 

Yaracuy, and Zulia; Map 292). In the park, found in sector El 

Mogote; 1,650–1,700 m.

This species forms large colonies in forest understory.

Isachne R. Br.

Isachne R. Br., Prodr. 196. 1810.

Annual or perennial plants; caespitose, sometimes rhizomatous or stoloniferous. Culms erect, decumbent or creeping, simple or branched. Sheaths rounded; ligules ciliate; blades 

flat, narrowly lanceolate to ovate. Inflorescences panicles, terminal, usually open. Spikelets obovoid to subglobose, biconvex, 2-flowered; disarticulation above or below the glumes; 

florets separated by a short internode and falling as a unit or 

glumes falling first; glumes subequal, nearly equal in length to 

spikelets, 5–7(–9)-nerved; lower florets staminate or bisexual, 

membranous to hardened, shorter than or equal to spikelets in 

length; paleas well-developed; upper florets pistillate or bisexual, 

± equal to spikelets in length; anthers 3. Caryopses ellipsoid to 

subglobose. [Micrairoideae.]

A tropical genus of ~100 species found in Mexico, Central 

America, the West Indies, South America (Colombia, Venezuela, 

the Guianas, Ecuador, Peru, Bolivia, and Brazil), Africa, Madagascar, Eurasia, Asia, and the Pacific Islands. Two or 3 species 

are found in Venezuela.

References. Hitchcock (1920); Sánchez-Ken et al. 

(2007); Webster et al. (1989).

Isachne is easily confused with Lasiacis (Griseb.) Hitchc., 

but the former has ciliate and the latter has membranous ligules.

Isachne rigens (Sw.) Trin., Gram. Panic. 252. 1826; Judziewicz, 

in Berry et al., Fl. Venez. Guayana 8: 135, fig. 92. 2004; 

Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos 

Venez. 2: 631–632, fig. 72. 2010. Panicum rigens Sw., 

Prodr. 23. 1788.

Isachne ligulata Swallen, Caldasia 2: 305. 1943.

Perennial plants. Culms 1–2(–10) m long, weak, trailing; flowering branches numerous, erect, 10–15(–30) cm tall, 

glabrous. Sheaths glabrous or puberulent; ligules 0.5(–2) mm 

long; blades narrowly oblong-lanceolate, (2–)5–6 cm × 1–5 mm, 

spreading, scabrous above and below. Inflorescences terminal, 

ovoid or oblong, 2–5(–10) cm long; branches and branchlets ascending or spreading. Spikelets 1.5–2 mm long; rachilla below 

the 2 florets minutely villous; glumes puberulent.

Found in Central America, the West Indies, and South 

America (Colombia, Venezuela, Guyana, Ecuador, and Peru). 

Widespread in Venezuela (Amazonas, Anzoátegui, Aragua, 

Bolívar, Distrito Federal, Lara, Mérida, Miranda, Monagas, 

Sucre, Táchira, and Trujillo; Map 293). In the park, fairly common in forest understory along trails and in roadside ditches; 

1,850–2,350(–3,000) m.

Lasiacis (Griseb.) Hitchc.

Lasiacis (Griseb.) Hitchc., Contr. U.S. Natl. Herb. 15: 16. 1910. Panicum

sect. Lasiacis Griseb., Fl. Brit. W.I. 551. 1864.

Perennial (rarely annual) plants, often bamboo-like (i.e., 

woody or lignified); caespitose. Culms erect, arching or scandent, simple or branched, herbaceous to woody; internodes solid 

or hollow; rooting at nodes. Sheaths open, rounded, margins free 

and overlapping; ligules membranous; blades linear to ovate, 

bases slightly to strongly asymmetrical. Inflorescences panicles, 

open or contracted; disarticulation below the glumes. Spikelets 

globose, obovoid or ellipsoid, with 2 florets. Glumes membranous, apices apiculate, lanate, usually shiny black when mature; 

lower glume shorter than spikelet, 5–13-nerved, bases saccate; 

upper glume ± equal in length to upper florets, 7–15-nerved, not 

saccate; lower floret sterile or staminate; lower lemma(s) membranous, apiculate, apices lanate; lower palea present (sometimes 

reduced); upper florets bisexual; upper lemma hardened, lanate; 

upper palea similar to lemma(s), but saccate below. Caryopses 

ovoid, obovoid or ± spherical. [Panicoideae.]

An American genus of 16 species found in North America 

(Greenland, Canada, and USA), Mexico, Central America, the 

MAP 293. Isachne rigens occurrence in Venezuela.
NUMBER 100 • 239

West Indies, and South America (all country-level political units). 

Twelve species occur in Venezuela.

References. Crins (1991); Davidse (1978); Morrone et al. (2012); Webster (1988); Webster et al. (1989).

KEY TO THE SPECIES OF LASIACIS

1a. Bases of leaf blades conspicuously cordate-auriculate; inflorescences 20–120 cm long . . . . . . . . . . . . . . . L. procerrima

1b. Bases of leaf blades not or slightly cordate-auriculate; inflorescences 5–12(–20) cm long . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Ligules of upper leaves readily visible, 1.5–3 mm long; branches of panicles reflexed or widely spreading; spikelets 

3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. ligulata

2b. Ligules of upper leaves not readily visible, <1.5 mm long; branches of panicles ascending to divergent; spikelets 

4–5(–6) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. nigra

Lasiacis ligulata Hitchc. & Chase, Contr. U.S. Natl. Herb. 18(7): 

337. 1917; Davidse, in Berry et al., Fl. Venez. Guayana 8: 

139, fig. 96. 2004.

Perennial, bamboo-like plants. Culms erect to arching, 

1–3(–10) m × 5–10 mm, woody, hollow; nodes glabrous. Sheaths 

glabrous to puberulent, margins ciliate; ligules 1.5–3 mm long, 

apices glabrous or ciliolate; blades narrowly to broadly lanceolate, 5–15 × 1–2.5 cm, bases asymmetrical, apices acuminate, 

glabrous or puberulent. Inflorescences 5–10(–20) cm long; 

branches reflexed or widely spreading; rachises glabrous or minutely scabrid. Spikelets obovoid, 3–4 mm long. Lower glume 

1–2 mm long, 7–11-nerved, glabrous with ciliate margins; upper 

glume 9–11-nerved; lemma(s) 9–11-nerved; palea shorter than 

or equal in length to florets. Caryopses ~2 mm long.

Found in Central America, the West Indies, and South 

America (Colombia, Venezuela, Trinidad and Tobago, the Guianas, Ecuador, Peru, Bolivia, Brazil, and Argentina). In Venezuela, recorded from Amazonas, Anzoátegui, Bolívar, Carabobo, 

Delta Amacuro, Miranda, Nueva Esparta, Sucre, and Trujillo 

(Map 294). In the park, found near the Laguna de Aguas Negras; 1,900 m.

Lasiacis nigra Davidse, Phytologia 29: 152. 1974; Davidse, in 

Berry et al., Fl. Venez. Guayana 8: 139, fig. 96. 2004.

Perennial, bamboo-like plants. Culms erect at base, arching 

above, 1–8 m × 3–5(–10) mm, woody, hollow; nodes glabrous. 

Sheaths pilose to glabrous, margins ciliate; ligules 0.5–1.5 mm 

long, glabrous or ciliate; blades linear to lanceolate, 5–10 × 0.5–

1.2(–2) cm, bases asymmetrical, slightly cordate-auriculate, 

and clasping the culm, margins scabrid, apices acuminate, pilose or glabrous. Inflorescences 5–12 cm long; branches ascending to diverging; rachises pilose to glabrous. Spikelets obovoid, 

4–5(–6) mm long, usually purple (when immature). Lower glume 

2–3 mm long, 5–13-nerved; upper glume 7–13-nerved; lemma 

9–11-nerved; palea shorter than or subequal in length to florets. 

Caryopses 2.5–3 mm long.

MAP 294. Lasiacis ligulata occurrence in Venezuela. MAP 295. Lasiacis nigra occurrence in Venezuela.
240 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Guyana, Ecuador, Peru, and Bolivia). In 

Venezuela, recorded from Aragua, Bolívar, Distrito Federal, Falcón, Lara, Mérida, Miranda, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, and Zulia (Map 295). In the park, found on both 

slopes of Guaramacal including near the Laguna de Aguas Negras and in Qda. Honda; 1,700–2,100 m.

Lasiacis procerrima (Hack.) Hitchc., Proc. Biol. Soc. Wash. 24: 

145. 1911; Davidse, in Berry et al., Fl. Venez. Guayana 8: 

139, fig. 98. 2004. Panicum procerrimum Hack., Oesterr. 

Bot. Z. 51: 431. 1901.

Annual or short-lived perennial plants; colonial, rarely solitary. Culms decumbent at base, simple or sparingly branched in 

the upper nodes, 1–4 m × 5–10 mm, herbaceous to woody, hollow; nodes glabrous; adventitious prop roots arising from lower 

nodes. Sheaths glabrous to puberulent and glaucous, auricles, if 

present, pubescent and ciliate; ligules 0.5–1.5 mm long, glabrous 

and minutely ciliolate; blades lanceolate, 15–30(–40) × 3–4(–6) 

cm, bases conspicuously cordate-auriculate, clasping the culm, 

apices acuminate, glabrous or minutely velutinous. Inflorescences highly branched and diffuse, 20–120 cm long and almost 

as wide; branches whorled at base; rachises scabrous. Spikelets obovoid, 2–3.5(–5) cm long. Lower glume 1.8–2 mm long, 

7–11-nerved; upper glume 9–11-nerved; lemma 9–11-nerved; 

palea shorter than florets. Caryopses ~2 mm long.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Guyana, Ecuador, Peru, and Brazil). 

In Venezuela, recorded from Amazonas, Anzoátegui, Aragua, 

Barinas, Bolívar, Carabobo, Cojedes, Distrito Federal, Falcón, 

Guárico, Lara, Mérida, Miranda, Portuguesa, Sucre, Táchira, 

Trujillo, Yaracuy, and Zulia (Map 296). In the park, found near 

La Divisoria de la Concepción; 1,300–1,500 m.

• Melinis P. Beauv.

Melinis P. Beauv., Ess. Agrostogr. 54. 1812.

Perennial or annual plants. Culms erect, ascending or decumbent, solid, rooting from lower nodes. Sheaths open; ligules 

membranous and ciliate; blades flat, filiform, linear or lanceolate, 

pubescent, viscid or not, aromatic or not. Inflorescences terminal panicles, lax; branches numerous, slender, and flexuous; disarticulation below the glumes, sometimes also below the florets. 

Spikelets laterally compressed, oblong, purplish with 2 florets; 

lower glume present and small or absent, unawned; upper glume 

equaling or exceeding florets in length, sometimes gibbous basally, 

5–7-nerved, awned or unawned; lower floret staminate or sterile; 

lower lemma almost the size of the upper glumes, not gibbous; 

upper floret bisexual, compressed; upper lemma glabrous, smooth, 

unawned; upper palea resembling upper lemma. [Panicoideae.]

A genus of ~20 species found in Africa and Madagascar, 

with several species introduced and naturalized in mostly tropical regions throughout the world. In the Americas, found in 

North America (USA), Mexico, Central America, the West Indies, and South America (Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina, and 

Uruguay). Two species occur in Venezuela.

References. Crins (1991); Morrone et al. (2012); 

Webster (1988); Webster et al. (1989); Zizka (1990).

• Melinis minutiflora P. Beauv., Ess. Agrostogr. 54, t. 11, fig. 4. 

1812; Judziewicz, in Berry et al., Fl. Venez. Guayana 8: 151, 

fig. 108. 2004; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 637–638, fig. 77. 2010.

Perennial plants; caespitose. Culms 0.4–1.9 m tall, ascending, branched, nodes bearded. Sheaths densely tomentose, viscid; 

ligules composed of hairs, 0.5–1.5 mm long; blades flat, linear, 

5–15 × 0.3–0.9 mm, densely viscid-pilose, aromatic. Inflorescences pyramidal, 5–20 cm long; pedicels shorter than spikelets. 

Spikelets 1.5–2.5 mm long, glabrous, usually purplish; lower 

glume absent or 0.3–0.5 mm long; upper glume 1.5–2.5 mm 

long, unawned; lower floret sterile; lower lemma 2-lobed, unawned or awned, awns (when present) 5–15 mm long; lower 

palea absent; upper lemma 1.5–2 mm long; upper palea slightly 

longer than upper lemmas. Caryopses ~1 mm long.

Native to tropical Africa; naturalized in North America 

(USA), Mexico, Central America, the West Indies, South America (Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, 

Bolivia, Brazil, Paraguay, Argentina, and Uruguay), Asia, Australia, and the Pacific Islands. In Venezuela, found in Amazonas, 

Aragua, Barinas, Bolívar, Carabobo, Cojedes, Distrito Federal, 

Guárico, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, and Zulia (Map 297). 

In the park, found in disturbed areas in Qda. Segovia and below 

Páramo del Pumar; 1,900–2,100 m.

MAP 296. Lasiacis procerrima occurrence in Venezuela.
NUMBER 100 • 241

Common Names. Capin melao and paja cebosa.

Although native to Africa, Melinis minutiflora was first described in 1812 from specimens collected in Brazil, where it certainly had arrived earlier (Parsons, 1972). In 1860, the species 

was brought from Brazil to Venezuela and was used for pasture 

in the Gran Sabana (Pittier, 1926: 402) from where it has spread. 

It is an aggressive invader in many parts of the world, including 

the Venezuelan Andes.

Ocellochloa Zuloaga & Morrone

Ocellochloa Zuloaga & Morrone, Syst. Bot. 34: 690. 2009.

Panicum sect. Stolonifera Hitchc. & Chase ex Pilg., in Engler, Nat. Pflanzenfam., 2nd ed., 14e: 16. 1940.

Perennial plants; stoloniferous or decumbent, rooting and 

branching at the lower nodes to erect or scandent. Culms hollow, 

rarely solid, simple or branching. Ligules membranous; blades 

flat, ovate-lanceolate to long-lanceolate, densely pilose to glabrous, shortly pseudopetiolate. Inflorescences panicles, pyramidal or oblong, composed of few to numerous, 1-sided racemose 

branches; branches arranged on either side of the axis and bearing secund, paired, short-pedicellate spikelets. Spikelets ellipsoid to lanceoloid, pilose to glabrous; shortly pedicellate. Lower 

glumes 3(–5)-nerved, shorter than spikelets; upper glumes shorter 

than or subequal to lower lemma, acute to acuminate apically, 

5(–7)-nerved; lower lemmas with or without 1 or 2(occasionally 3) pairs of crateriform, ocellate glands; lower paleas hyaline, 

glabrous; staminate flowers present or absent (bisexual flowers 

rarely present); upper anthecia ellipsoid to lanceoloid, glabrous, 

smooth and shiny, hardened, shortly stipitate at the base; upper 

lemmas with margins inrolled over the paleas. [Panicoideae.]

An American genus of 12 species found in Mexico, the 

Lesser Antilles, Central America, and South America (Colombia, 

Venezuela, Ecuador, Peru, Bolivia, Brazil, and Argentina). Four 

species occur in Venezuela.

References. Morrone et al. (2012); Sede et al. 

(2009); Zuloaga and Sendulsky (1988).

Ocellochloa pulchella (Raddi) Zuloaga & Morrone, in Sede et 

al., Syst. Bot. 34: 690. 2009. Panicum pulchellum Raddi, 

Agrostogr. Bras. 42. 1823; Dorr et al., Contr. U.S. Natl. 

Herb. 40: 57. 2000 [2001]; Zuolaga, in Berry et al., Fl. 

Venez. Guayana 8: 188. 2004.

Perennial (?) plants. Culms decumbent, creeping and rooting 

at the nodes, 10–65 cm × 2–3 mm, nodes densely pilose, internodes 2–8 cm long. Sheaths shorter than internodes, pilose to 

glabrous, margins ciliate; ligules minute, membranous, ciliate; 

blades ovate-lanceolate, 1.5–7 × 0.5–2 cm, bases asymmetrical 

and cordate, margins long-ciliate at bases, apices acuminate, strigose to glabrous above and below. Inflorescences terminal solitary 

panicles, 5–10(–18) × 1–2.5(–4) cm, composed of 5–20 branches, 

ascending or reflexed, divergent from the axis; branches 3-angled, 

flattened on 1 side, densely hispid to scabrous; spikelets solitary 

or paired, 1 subsessile (often abortive), 1 short-pedicellate. Spikelets narrowly ellipsoid, ~2 mm long; upper glumes and lower lemmas subequal (or the upper glumes shorter); lower glumes ovate, 

3-nerved; upper glumes 5-nerved; lower lemmas 5-nerved, bearing 2 crateriform or ocellate glands; lower paleas hyaline, margins short-ciliate, otherwise glabrous; staminate flowers usually 

absent. Caryopses ellipsoid, ~1 mm long.

Found in Mexico, Central America, the Lesser Antilles (Martinique), and South America (Colombia, Venezuela, Ecuador, 

MAP 297. Melinis minutiflora occurrence in Venezuela.

MAP 298. Ocellochloa pulchella occurrence in Venezuela.
242 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Peru, Bolivia, Brazil, and Argentina). Found almost throughout 

Venezuela (Amazonas, Anzoátegui, Apure, Aragua, Barinas, 

Carabobo, Cojedes, Distrito Federal, Guárico, Lara, Mérida, 

Miranda, Monagas, Portuguesa, Sucre, Táchira, and Yaracuy; 

Map 298). In Guaramacal, recorded from the understory of and 

openings in forest near sector El Paramito; 1,600 m.

This species is remarkable for the presence of 1 or 2 (rarely 3) 

pairs of microscopic (but visible at 10× magnification) crateriform 

or ocellate glands on the outer surfaces of the upper lemmas.

Paspalum L.

Paspalum L., Syst. Nat., ed. 10, 846, 855, 1359. 1759.

Reimarochloa Hitchc., Contr. U.S. Natl. Herb. 12(6): 198. 1909.

Thrasyopsis Parodi, Bol. Soc. Argent. Bot. 1: 293. 1946.

Annual or perennial plants; caespitose, rhizomatous or 

stoloniferous. Culms erect, spreading or prostrate. Sheaths 

open; ligules membranous, glabrous or ciliate; blades mostly 

flat, linear, lanceolate or linear-lanceolate. Inflorescences terminal, sometimes also axillary, panicles, with 1 to many spikelike 

racemes; raceme axes flattened, usually narrowly to broadly 

winged, usually terminating in a spikelet; disarticulation below 

the glumes. Spikelets subsessile to shortly pedicellate, dorsally 

compressed, solitary or paired, in 2 rows along 1 side of the racemes, with 2 florets; lower florets sterile; upper florets sessile or 

stipitate, bisexual; lower glumes absent or rudimentary and present only in some spikelets, (0)1-nerved, unawned; upper glumes 

and lower lemmas subequal, membranous, unarmed; lower paleas absent or rudimentary; upper lemmas hardened, involute, 

clasping the paleas; upper paleas hardened. Anthers 2. Stigmas 

2, plumose. Caryopses orbicular to ellipsoid, plano-convex or 

flattened. [Panicoideae.]

A genus of ~350 species found in the Americas, Europe, Africa, Madagascar, Asia, Australia, and the Pacific Islands. The 

majority of species are American, found in North America (Canada and USA), Mexico, Central America, the West Indies, and 

South America (all country-level political units). Approximately 

100 species occur in Venezuela.

References. Chase (1929); Crins (1991); Denham 

(2005); Morrone et al. (1995, 2012); Rodríguez (2006); Rua et 

al. (2010); Webster (1988); Webster et al. (1989).

In a combined molecular and morphological study Rua et 

al. (2010) found Paspalum to be generally monophyletic, but 

three of the four currently recognized subgenera and many of the 

informal morphological groupings were polyphyletic. The only 

morphological synapomorphy for the genus appears to be the 

distinctive plano-convex spikelets.

Spikelet orientation is the only morphological character that 

separates Panicum from Axonopus. In the former genus fertile 

lemmas and upper glumes are turned toward the rachis.

KEY TO THE SPECIES OF PASPALUM

1a. Inflorescences with 60–200 racemes; leaf blades >50 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. densum

1b. Inflorescences with <50 racemes; leaf blades <35 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Racemes 2(3) per inflorescence; spikelets solitary (rarely paired), sparingly puberulent . . . . . . . . . . . . . P. distichum

2b. Racemes 2–50 per inflorescence; spikelets solitary or paired, pubescent or glabrous . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Spikelets solitary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. penicillatum

 3b. Spikelets paired . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 4a. Spikelets pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 5a. Ligules 2–3 mm long; bases of leaf blades auriculate . . . . . . . . . . . . . . . . . . . . . . . . . P. macrophyllum

 5b. Ligules ~0.5 mm long; bases of leaf blades not auriculate . . . . . . . . . . . . . . . . . . . . . . . P. paniculatum

 4b. Spikelets glabrous or sparingly pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. inconstans

Paspalum densum Poir., in Lamarck, Encycl. 5: 32. 1804; Zuloaga and Judziewicz, in Berry et al., Fl. Venez. Guayana 8: 

223, fig. 182. 2004.

Robust perennial plants; caespitose. Culms 100–300 cm tall, 

erect; nodes glabrous to densely pubescent. Sheaths keeled, glabrous or ciliate distally; ligules 1.5–3 mm long; blades flat or plicate, linear, 50–80(–100) × 1–2 cm, bases not auriculate, sparsely 

to densely long-hirsute. Inflorescences terminal, 12–30(–40) cm 

long, with 60–200 racemes; racemes 4–9 cm long. Spikelets orbicular to orbicular-obovoid, 1.5–2.5 mm long, paired, glabrous; 

upper glumes and lower lemmas membranous, 3-nerved.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Trinidad and Tobago, the 

Guianas, Bolivia, and Brazil). In Venezuela, reported from Amazonas, Anzoátegui, Apure, Barinas, Bolívar, Carabobo, Cojedes, 

Guárico, Mérida, Monagas, Portuguesa, Trujillo, and Zulia 

(Map 299). Found in disturbed areas in El Campamento below 

Cerro El Diablo; 1,800–2,000 m.

Paspalum distichum L., Syst. Nat., ed. 10, 855. 1759.

Perennial plants; rhizomatous or caespitose. Culms 30–

45(–65) cm tall, erect or subdecumbent; nodes glabrous. Sheaths 

keeled, glabrous to sparsely long-pubescent; ligules 1–2 mm long; 

blades flat or involute, linear-lanceolate, 13–15 × 0.5–0.6 cm, 

bases not auriculate, glabrous or pubescent. Inflorescences terminal, <6.5 cm long, with 2(3) racemes; racemes 1.5–7 cm long. 

Spikelets ellipsoid, ~2.5–3 mm long, solitary (rarely paired), 

sparingly puberulent; lower glumes absent or, if present, ~1 mm 

long; upper glumes 3-nerved, sparsely pubescent; lower lemmas 

3-nerved, glabrous.
NUMBER 100 • 243

A pantropical species found in North America (USA), 

Mexico, the West Indies, and South America (all country-level 

political units); adventive in Europe. In Venezuela, known from 

Apure, Aragua, Carabobo, Delta Amacuro, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Sucre, Táchira, Trujillo, 

and Yaracuy (Map 300). In the park, found near the Laguna de 

los Cedros; 1,900 m.

Paspalum inconstans Chase, Contr. U.S. Natl. Herb. 24(8): 446. 

1927.

Perennial plants; caespitose. Culms 20–100 cm tall, ascending; nodes glabrous. Sheaths pilose; ligules 4 mm long; blades 

linear to linear-lanceolate, 5–25 × 0.5–1 cm, bases not auriculate, pubescent. Inflorescences terminal and axillary, 10–20 cm 

long, with 2 or 3(–8) racemes; racemes 5–8 cm long. Spikelets 

ovoid or ellipsoid, 2–2.5 mm long, paired, glabrous or sparingly 

pubescent; upper glumes membranous, shorter than the spikelets, 5-nerved, glabrous or sparsely pubescent; lower lemmas 

equaling the spikelets, 5-nerved.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia). In Venezuela, found only in the Andes (Lara, 

Mérida, Táchira, and Trujillo; Map 301). In the park, found in 

various localities on both slopes of Guaramacal; 1,900–2,150 m.

Paspalum macrophyllum Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 

1: 76. 1815 [1816]; ibid. [qu.] 1: 92. 1815 [1816].

Perennial plants; caespitose. Culms 20–45(–150) cm tall, 

ascending; nodes glabrous. Sheaths papillose-pilose or glabrous; 

ligules 2–3 mm long; blades flat, lanceolate, 5–15(–25) × 1–2 cm, 

bases auriculate, glabrous or finely pubescent below. Inflorescences terminal, 10–15 cm long, with 5–12 racemes; racemes 3–5 

cm long. Spikelets ellipsoid to obovoid, 2–3 mm long, paired, 

puberulent; upper glumes 5-nerved, finely pubescent; lower lemmas 3–5-nerved, glabrous or appressed pubescent.

Found in South America (Colombia, Venezuela, Ecuador, 

Peru, and Bolivia); introduced in North America (USA), the 

Greater Antilles (Puerto Rico), and the Pacific Islands. In Venezuela, the species has been collected in the Andes (Lara, Mérida, 

Táchira, and Trujillo) and in the Cordillera de la Costa (Aragua, 

MAP 299. Paspalum densum occurrence in Venezuela.

MAP 300. Paspalum distichum occurrence in Venezuela.

MAP 301. Paspalum inconstans occurrence in Venezuela.
244 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Falcón, and Yaracuy; Map 302). Our records are from various localities in the park on both slopes of Guaramacal; 1,850–2,000 m.

Paspalum paniculatum L., Syst. Nat., ed. 10, 855. 1759; Zuloaga and Judziewicz, in Berry et al., Fl. Venez. Guayana 8: 

224. 2004.

Perennial plants; caespitose, sometimes stoloniferous. 

Culms 30–100 cm tall, erect; nodes glabrous to pubescent. 

Sheaths keeled, papillose-hispid; ligules ~0.5 mm long; blades 

flat, lanceolate to linear-lanceolate, 8–35 × 0.7–1(–3) cm, bases 

not auriculate, scabrous to glabrescent. Inflorescences terminal, 

5–30 cm long, with (5–)20–50 racemes; racemes 1–5(–10) cm 

long. Spikelets orbicular or narrowly obovate, 1–1.5 mm long, 

paired, pubescent; lower glumes absent; upper glumes membranous, 3-nerved, pubescent; lower lemmas similar to glumes but 

subglabrous in the middle.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Trinidad and Tobago, 

the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina, and Uruguay); introduced in North America (USA), Africa, 

Australia, and the Pacific Islands. In Venezuela, recorded from 

Anzoátegui, Apure, Aragua, Barinas, Bolívar, Carabobo, Delta 

Amacuro, Distrito Federal, Lara, Mérida, Miranda, Monagas, 

Nueva Esparta, Portuguesa, Táchira, Trujillo, Yaracuy, and 

Zulia (Map 303). Our records are from various localities in the 

park on both slopes of Guaramacal; 1,900–2,000 m.

This species is considered to be invasive in many parts of 

its range.

Paspalum penicillatum Hook. f., Trans. Linn. Soc. London 20: 

171. 1847.

Paspalum prostratum Scribn. & Merr., Bull. Div. Agrostol., U.S.D.A. 24: 9. 

[9 Jan] 1901, non Nash, [Oct] 1901; Dorr et al., Contr. U.S. Natl. Herb. 

40: 58. 2000 [2001]; Briceño, in Morillo et al., Bot. Ecol. Monocot. 

Páramos Venez. 2: 658, fig. 9. 2010.

Annual plants; caespitose. Culms (15–)30–40(–150) cm tall, 

erect or ascending; nodes glabrous to sparingly pilose. Sheaths 

keeled, glabrous or puberulent, margins ciliate; ligules minute, 

to 0.5 mm long; blades linear, lanceolate or linear-lanceolate, 

4–5(–20) × 0.4–0.5(–1.5) cm, bases not auriculate, pilose. Inflorescences terminal, 5–20 cm long, with 3–9(–15) racemes; 

MAP 302. Paspalum macrophyllum occurrence in Venezuela.

MAP 303. Paspalum paniculatum occurrence in Venezuela. MAP 304. Paspalum penicillatum occurrence in Venezuela.
NUMBER 100 • 245

racemes 1–4 cm long. Spikelets ellipsoid to obovoid-ellipsoid, 

solitary, glabrous; lower glumes absent; upper glumes and lower 

lemmas membranous, as long as spikelets, 3-nerved.

Found in Mexico and South America (Colombia, Venezuela, 

Ecuador, Peru, and Bolivia). In Venezuela, this species is recorded 

from the Andes (Lara, Mérida, Táchira, and Trujillo) and the 

Cordillera de la Costa (Aragua, Distrito Federal, and Guárico; 

Map 304). In the park, found in subpáramo in the Qda. Jirajara; 

2,100–2,800 m.

Morrone et al. (1995) recognized Paspalum penicillatum

and P. prostratum as distinct species. The two differ only slightly; 

there are minor size differences with respect to the width of rachises and dimensions of spikelets. We follow Rodríguez (2006) 

in considering these names to be synonyms.

• Poa L.

Poa L., Sp. Pl. 67. 1753.

Perennial or rarely annual plants; caespitose or rhizomatous. 

Culms unbranched. Sheaths split part of the way to base; ligules 

truncate, glabrous or shortly pilose; blades flat or involute, linear, 

apices navicular, glabrous, pubescent or scabrous. Inflorescences 

terminal panicles, lax or contracted; flowers bisexual or unisexual (and then the plants dioecious). Spikelets pedicellate, laterally 

compressed, oblong, ovate or elliptic, (1–)2–6-flowered; disarticulation above the glumes and below the florets. Glumes 2, subequal, 

elliptic, shorter than spikelets, acute; lower glumes 1-nerved; 

upper glumes 3-nerved; lemmas generally carinate, acute or blunt, 

awnless, usually 5(–7)-nerved, membranous; midnerve, marginal 

and intermediate nerves often pubescent; paleas generally shorter 

than lemmas; lodicules 2; anthers 3; styles 2, separate, short or 

obsolete. Caryopses subterete, ellipsoid. [Pooideae.]

A cosmopolitan genus of ~500 species found at high elevations in the subtropics and tropics and widespread in the Americas: North America (Greenland, Canada, and USA), Mexico, 

Central America, the West Indies, and South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Chile, Paraguay, 

Argentina, and Uruguay). Seven species occur in Venezuela, 

mainly in the Andes and in the Cordillera de la Costa.

References. Gillespie et al. (2007); Tucker (1996).

Poa is related to Festuca, from which it can be distinguished 

by its awnless anthers and navicular leaf apices.

• Poa annua L., Sp. Pl. 68. 1753; Briceño, in Morillo et al., Bot. 

Ecol. Monocot. Páramos Venez. 2: 666, fig. 98. 2010.

Annual or short-lived perennial plants; caespitose. Culms 

erect, spreading or prostrate, sometimes rooting at the nodes, 

5–35 cm tall, glabrous. Ligules membranous, 2–5 mm long; 

blades flat, 5–10 × 0.1–0.3 cm, margins minutely scabrous. Inflorescences solitary, ovate to pyramidal, 1.5–12 cm tall; branches 

ascending to spreading, glabrous. Spikelets ovate or oblong, (1.5–)

3–5(–8) mm long, laterally compressed, 3–10-flowered; rachillae 

often visible. Glumes persistent, unequal, keeled, glabrous; lower 

glumes 1.5–2(–3) mm long; upper glumes 2–2.5(–4) mm long; 

lemmas elliptic to oblong, 2.5–4 mm long, keeled, 5-nerved, sparingly to densely pubescent on the nerves or glabrous.

Native to Europe and now cosmopolitan. In the Americas, 

found in North America (Greenland, Canada, and USA), Mexico, Central America, the West Indies, and South America (Colombia, Venezuela, Guyana, Ecuador, Peru, Bolivia, Brazil, Chile, 

Paraguay, Argentina, and Uruguay). In Venezuela, reported from 

Apure, Aragua, Barinas, Bolívar, Distrito Federal, Lara, Mérida, 

Miranda, Táchira, Trujillo, and Zulia (Map 305). In the park, 

found on both slopes and in the Páramo de Guaramacal; (1,600–)

1,900–3,100 m.

This species might have the widest geographical distribution of any species of angiosperm as it is found in the Americas, 

Africa, Asia, Australia, and Antarctica.

Polypogon Desf.

Polypogon Desf., Fl. Atlant. 1: 66. 1798 [1800].

Chaetotropis Kunth, Énum. Gramin. 1: 72. 1829.

Perennial or annual plants; caespitose or rhizomatous. 

Culms erect or decumbent; rooting at lower nodes. Sheaths 

smooth to scabridulous; ligules membranous, erose, ciliate; 

blades linear, flat. Inflorescences terminal panicles or spikelike, 

contracted to dense; branches, rachises, and pedicels scabrous. 

Spikelets ellipsoid, laterally compressed, 1-flowered; disarticulation slightly below glumes (spikelets stipitate, falling intact); 

florets perfect; glumes 2, equal or subequal, broadly lanceolate, longer than spikelets, entire or sometimes 2-lobed, awned, 

1-nerved; lemmas lanceolate, much shorter than glumes, hyaline, 

obscurely 1–3(–5)-nerved, 2-lobed or 2-aristate, awned apically; 

paleas elliptic, slightly shorter than or equaling lemmas, hyaline, 

MAP 305. Poa annua occurrence in Venezuela.
246 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

2-nerved, apices 2-dentate; lodicules 2; anthers 1–3. Caryopses 

slightly compressed, ellipsoid to obovoid. [Pooideae.]

A cosmopolitan genus of ~18 species occurring in warm 

temperate regions and mountainous areas in the tropics. In the 

Americas, native and introduced species found in North America 

(Canada and USA), Mexico, Central America, the Greater Antilles, and South America (Colombia, Venezuela, Ecuador, Peru, 

Bolivia, Brazil, Chile, Paraguay, Argentina, and Uruguay). Three 

species occur in Venezuela.

References. Giraldo-Cañas (2004); Tucker (1996).

Polypogon is closely related to Agrostis, from which it can 

be distinguished by its spikelets that disarticulate below the 

glumes, often at the base of a stipe.

Polypogon elongatus Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 

110. 1815 [29 Jan 1816]; ibid. [qu.] 1: 134. 1815 [29 Jan 

1816], non Lag., [Jun-Dec] 1816 (“elongatum”); Briceño, 

in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 

675–676, fig. 105. 2010. Chaetotropis elongata (Kunth) 

Björkman, Symb. Bot. Upsal. 17: 14. 1960.

Perennial plants; caespitose. Culms 50–100 cm tall, erect or 

decumbent at base, unbranched. Sheaths glabrous; ligules nearly 

glabrous, 3.5–8 mm long, lacerate apically, decurrent on sheath 

margins; blades 10–25(–30) × 0.4–0.8(–1.5) cm, scaberulous 

above and along margins, glabrous or scaberulous below. Panicles 10–25(–30) cm long; branches erect or nodding, somewhat 

loose and interrupted, especially below; spikelets pedicellate, 

stipes 1.5–2.5 mm long, deciduous. Glumes subequal, lanceolate, lower and upper glumes 3–5 mm long, each with an awn 

1–2(–4) mm long; lemmas glabrous, awns 1–2.5(–3) mm long; 

paleas shorter than lemmas.

Found in Mexico, Central America, and South America 

(Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Chile, Argentina, and Uruguay); adventive in North America (USA). In 

Venezuela, found in Apure, Aragua, Barinas, Bolívar, Carabobo, 

Distrito Federal, Lara, Mérida, Miranda, Táchira, Trujillo, and 

Zulia (Map 306). In the park, found in open areas and along forest edges near the Laguna de Aguas Negras and on both slopes of 

Guaramacal; 1,800–3,100 m.

Rugoloa Zuloaga

Rugoloa Zuloaga, in J. M. Acosta et al., Pl. Syst. Evol. [epublished 29 Mar 

2014].

Perennial plants; rhizomatous. Culms erect or decumbent, 

rooting and branching at the lower nodes. Sheaths open; ligules membranous, ciliate; blades ovate-lanceolate to lanceolate, 

flat or inrolled. Inflorescence a terminal panicle. Disarticulation 

below the spikelets, which fall entire from the pedicels. Spikelets 2-flowered, plump, terete, laterally compressed, pilose to 

glabrous. Lower glume less than 1⁄2 the length of the spikelet, 

membranous, 3-nerved; upper glume and lower lemma subequal, ± equal to spikelet, membranous, 5-nerved; lower palea 

well-developed or absent; upper (fertile) floret bisexual, lemma 

indurate, awnless. Lodicules 2. Anthers usually 3. Stigmas 2, plumose. Caryopses ellipsoid. [Panicoideae.]

An American genus of three species found in Mexico, Central America, the West Indies, and South America (Colombia, 

Venezuela, Trinidad and Tobago, the Guianas, Ecuador, Peru, 

Bolivia, Brazil, Paraguay, and Argentina). All three species are 

found in Venezuela.

References. Acosta et al. (2014); Aliscioni et al. 

(2003); Crins (1991); Hitchcock and Chase (1910, 1915); Morrone et al. (2012); Webster (1988); Webster et al. (1989); Zuloaga et al. (1992).

Molecular phylogenetic data led Acosta et al. (2014) to recognize this small genus of species formerly placed in Panicum s.l. 

Acosta et al. (2014) also observed that the genus is characterized 

by a number of morphological characters: inflorescences with 

lax panicles; spikelets ± crowded and unilaterally disposed on 

the branches or in short branchlets; spikelets narrowly ellipsoid, 

hispid with caducous hairs or glabrous; upper anthecium ellipsoid, indurate, with stomata, simple papillae, and prickle hairs 

only present toward the apex; and silica bodies occasionally 

present at the tip of the lemma.

Rugoloa polygonata (Schrad.) Zuloaga, in J. M. Acosta et al., 

Pl. Syst. Evol. [epublished 29 Mar 2014]. Panicum polygonatum Schrad., in Schultes, Mant. 2: 256. 1824; Zuolaga, 

in Berry et al., Fl. Venez. Guayana 8: 188, fig. 154. 2004. 

Dallwatsonia polygonata (Schrad.) J. R. Grande, Phytoneuron 2014–22: 4. 2014.

Perennial plants; shortly rhizomatous. Culms 30–100 cm 

tall, decumbent and rooting at lower nodes, becoming erect; 

nodes densely pilose. Sheaths glabrous or rarely papillose-pilose; 

MAP 306. Polypogon elongatus occurrence in Venezuela.
NUMBER 100 • 247

ligules membranous, ciliate; blades lanceolate, 5–15(–20) cm × 

5–15 mm, bases cordate, apices attenuate, glabrous or sparsely 

pilose above and below. Inflorescences lax, pyramidal panicles, 

9–25 × 3–15 cm; primary branches ascending or spreading, 

becoming shorter toward apex. Spikelets narrowly ellipsoid, 

1.3–1.6 mm long, pubescent or glabrous. Glumes similar, ovate; 

lower glume ~1 mm long, 3-nerved; upper glume ~1.4 mm long, 

3- or 5-nerved. Lower lemma ~1.4 mm long, 3–5-nerved. Lower 

palea ~1 mm long, hyaline, glabrous or absent. Caryopses ~1 × 

0.5 mm.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, Trinidad and Tobago, 

Guyana, French Guiana, Ecuador, Peru, Bolivia, Brazil, Paraguay, and Argentina). Widespread in Venezuela (Amazonas, 

Apure, Aragua, Bolívar, Carabobo, Distrito Federal, Falcón, 

Lara, Mérida, Miranda, Monagas, Portuguesa, Táchira, and 

Zulia; Map 307). In Guaramacal, almost always in or near coffee plantations such as those found near La Divisoria de la Concepción; ~1,500 m.

Zuloaga et al. (1992) placed this species in Panicum subgen. Phanopyrum (Raf.) Pilg., but subsequent analyses (Aliscioni 

et al., 2003) indicate that subgen. Phanopyrum is polyphyletic 

and that a monophyletic Phanopyrum (Raf.) Nash can only 

be recognized if it is monotypic and restricted to Phanopyrum 

gymnocarpon (Elliott) Nash. Aliscioni et al. (2003) left Panicum 

polygonatum as incertae sedis pending further investigation. 

Subsequently, Acosta et al. (2014), in a review of Paspaleae J. 

Presl subtribe Otachyriinae Butzin, placed Panicum polygonatum and two other species of Panicum in the genus Rugoloa, 

which they recognized as a strongly supported clade that had no 

apparent relationships to other genera in the Otachyriinae.

• Setaria P. Beauv.

Setaria P. Beauv., Ess. Agrostogr. 51, 178. 1812, nom. cons.

Annual or perennial plants; caespitose or rarely rhizomatous. Culms erect or decumbent. Sheaths rounded to carinate; 

ligules membranous and ciliate or composed of hairs; blades flat, 

folded or involute, linear to lanceolate, sometimes plicate or narrowed to a pseudopetiole. Inflorescences terminal panicles, dense 

and spikelike or loose and open; disarticulation usually below 

the glumes, spikelets falling intact, bristles persistent. Spikelets 

usually lanceoloid-ellipsoid, rarely globose, subsessile to shortpedicellate, in fascicles or single on short branches, some or all 

subtended by 1 to several bristles. Lower glumes membranous, 

less than 1⁄2 the length of the spikelets, 1–7-nerved; upper glumes 

membranous to herbaceous, 1⁄2 as long to nearly equaling the 

upper lemmas in length, 3–9-nerved; lower florets staminate or 

sterile; lower lemmas membranous, equaling the upper lemmas in 

length, rarely absent, 5–7-nerved; lower paleas hyaline to membranous, rarely absent or reduced; upper florets bisexual; upper 

lemmas and paleas indurate, transversely rugose; anthers 3; styles 

2. Caryopses ellipsoid to subglobose, compressed. [Panicoideae.]

A genus of 100–125 species found in temperate and tropical areas worldwide. In the Americas, found in North America 

(Canada and USA), Mexico, Central America, the West Indies, 

and South America (Colombia, Venezuela, Curaçao, Trinidad 

and Tobago, the Guianas, Ecuador, Peru, Bolivia, Brazil, Chile, 

Paraguay, Argentina, and Uruguay). Seventeen species occur in 

Venezuela.

References. Crins (1991); Kellogg et al. (2009); 

Morrone et al. (2012); Pensiero (1999); Rominger (1962); Webster (1988); Webster et al. (1989).

The genus as currently construed apparently is not monophyletic and likely will be split into several genera (Kellogg et al., 

2009). Nonetheless, the same study that refuted its monophyly 

identified several large, geographically coherent clades.

• Setaria parviflora (Poir.) Kerguélen, Lejeunia, n.s., 120: 161. 

1987; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 681–682, fig. 111. 2010. Cenchrus parviflorus Poir., in Lamarck, Encycl. 6: 52. 1804.

Setaria geniculata P. Beauv., Ess. Agrostogr. 51, 169, 178. 1812.

The nominate variety is the only one known from Guaramacal and Venezuela.

• Setaria parviflora var. parviflora

Perennial plants; caespitose or rhizomatous, rhizomes short. 

Culms 30–120 cm tall, erect or sometimes decumbent, branching. Sheaths glabrous; ligules <1 mm long, composed of stiff 

hairs; blades linear to lanceolate 10–25 cm × 2–10 mm, glabrous or with a few long hairs above. Peduncles 5–32 cm long, 

glabrous, scabrid at the apex. Inflorescences dense, spikelike 

panicles, 1–8 × 2.5 cm, uniform width throughout their length; 

rachises minutely pubescent; bristles 4–12, 2–10 mm long, yellow or bronze, scabrous. Spikelets 2–3 mm long, elliptical. Lower 

MAP 307. Rugoloa polygonata occurrence in Venezuela.
248 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

glumes ~1 mm long, 3(–4)-nerved; upper glumes 1–1.5 mm long, 

5-nerved; lower florets often staminate; lower lemmas ± equal to 

spikelet, 5–7-nerved; lower paleas equaling lower lemmas; upper 

lemmas transversely rugose. Caryopses ellipsoid.

Evidently native to Africa; adventive in North America 

(Canada and USA), Mexico, Central America, the West Indies, 

South America (Colombia, Venezuela, Trinidad and Tobago, the 

Guianas, Ecuador, Peru, Bolivia, Brazil, Chile, Paraguay, Argentina, and Uruguay), Europe, Asia, Australia, and the Pacific Islands. Widespread in Venezuela (Amazonas, Anzoátegui, Apure, 

Aragua, Barinas, Bolívar, Carabobo, Cojedes, Delta Amacuro, 

Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, 

Monagas, Nueva Esparta, Portuguesa, Táchira, Trujillo, Yaracuy, and Zulia; Map 308). In the park, collected on the path to 

La Cueva below Cerro El Diablo; 1,800–2,000 m.

This species was long thought to be indigenous to South 

America, but Kellogg et al. (2009) determined that it is part of 

a strongly supported clade of African species, and it appears to 

have been introduced into the Americas.

Sporobolus R. Br.

Sporobolus R. Br., Prodr. 169. 1810.

Annual or perennial plants; usually caespitose, sometimes 

rhizomatous, rarely stoloniferous. Culms erect, glabrous. Sheaths 

open, glabrous, apices ciliate or ciliolate; ligules ciliate; blades 

flat, folded, involute, sometimes terete, pubescent or glabrous. 

Inflorescences terminal panicles, open or contracted, rarely 

spikelike. Spikelets bisexual, rounded to laterally compressed, 

1(–3)-florets per spikelet; disarticulation above the glumes or 

remaining intact until extrusion of the seed. Glumes equal or 

subequal, usually shorter than floret, 1-nerved or nerveless, 

membranous, or spikelet remaining intact; lemmas 1(3)-nerved, 

usually acute, membranous or chartaceous, unawned; paleas 

equal to or shorter than lemma, glabrous, 2-nerved; anthers 2(3). 

Caryopses utricle-like, with the mature pericarp usually gelatinous when wetted and extruding the seed, ovoid, ellipsoid, fusiform or quadrangular. [Chloridoideae.]

A genus of ~160 species occurring in tropical and subtropical regions worldwide. In the Americas, found in North America 

(Canada and USA), Mexico, Central America, the West Indies, 

and South America (all country-level political units). Nine species are found in Venezuela, some of which are aggressive weeds 

in cropland.

References. Clayton (1965); Giraldo-Cañas and 

Peterson (2009); Ortiz-Diaz and Culham (2000); Peterson et al. 

(1997); Shrestha et al. (2004).

Sporobolus indicus (L.) R. Br., Prodr. 170. 1810; Briceño, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 683–684, 

fig. 112. 2010. Agrostis indica L., Sp. Pl. 63. 1753.

Perennial plants; densely caespitose. Culms 50–90 cm 

tall. Sheaths keeled below, 5–15 cm long, glabrous, apices ciliolate; ligules minute, ciliolate; blades flat, lanceolate, 

10–35 cm × 3–7 mm, glabrous above and below. Inflorescences 

solitary, dense spikelike panicles, 10–30 cm × 5–15 mm. Spikelets ~2–3 mm long. Glumes subequal; lemmas ovate, 2–2.5 mm 

long, acute or obtuse, 1-nerved, glabrous; paleas similar to lemmas, 1.5–2.2 mm long, faintly 2-nerved, glabrous; anthers 3. 

Caryopses quadrangular, laterally compressed, ~1 mm long.

Found in North America (USA), Mexico, Central America, 

the West Indies, and South America (Colombia, Venezuela, 

MAP 308. Setaria parviflora var. parviflora occurrence in Venezuela.

MAP 309. Sporobolus indicus occurrence in Venezuela.
NUMBER 100 • 249

Trinidad and Tobago, Ecuador, Peru, Bolivia, Brazil, Chile, 

Paraguay, Argentina, and Uruguay). Widespread in Venezuela 

(Apure, Aragua, Barinas, Bolívar, Carabobo, Distrito Federal, 

Falcón, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Táchira, and Trujillo; Map 309). In the park, collected 

in open wet areas near the Laguna de Aguas Negras and between 

Qda. Honda and Pele El Ojo; 1,800–2,100 m.

Sporobolus indicus s.l. is polymorphic, and as currently 

construed, it is a complex of 11 species found throughout the 

tropics and subtropics (Clayton, 1965; Ortiz-Diaz and Culham, 

2000; Shrestha et al., 2004). Sporobolus indicus s. str. could be 

confused with S. jacquemontii Kunth, which also belongs to this 

complex and also occurs in Venezuela, but our species has solid 

internodes and a ± dense inflorescence (vs. hollow internodes 

and a ± lax inflorescence). In addition, S. jacquemontii generally 

is found at lower elevations.

Smilacaceae

L. J. DORR AND B. STERGIOS

Lianas (our species), shrubs or herbs; herbaceous or suffruticose; perennial or annual; rhizomatous; rhizomes tuberculate 

or elongate; roots filiform. Stems terete or angular, unarmed or 

armed with prickles; terminal branches straight, flexuous or zigzag; lateral shoots with 1 or 2 adaxial scales. Leaves simple, alternate, distichous or decussate; leaf blades linear, oblong, ovate 

or lanceolate (sometimes reduced to scales), apices acuminate 

to acute, margins entire or armed with prickles, bases acute or 

cordate, 3–9-veined from the base, secondary veins reticulate; 

petiolate, petioles typically forming a sheath that terminates in 

a pair of tendrils. Inflorescences umbellate (a pseudoumbellate 

cyme), solitary or aggregated in racemes, axillary, pedunculate; 

peduncles ending in a thickened receptacle. Flowers actinomorphic, unisexual (the plants dioecious), 3-merous, pedicellate; tepals 6, free or connate; staminate flowers: 6 free stamens in 2(3) 

whorls, anthers 1- or 2-thecal, basifixed, pistillode sometimes 

present; pistillate flowers: 0–6 staminodes; ovary superior, globose, 3-locular; ovules 1 or 2 per locule; styles short or absent; 

stigmas 3. Fruit a globose berry, orange, red, blue or black; seeds 

1–3, rounded to ovoid, reddish, orange, brown or black.

As construed here the family includes only Smilax L., a large 

pantropical and temperate genus of 200–300 species. The family 

is present throughout the Americas, including North America 

(Canada and USA), Mexico, Central America, the West Indies, 

and South America (Colombia, Venezuela, Trinidad and Tobago, 

the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina, 

and Uruguay). Approximately 100 species of Smilax are found in 

the neotropics, including ~14 in Venezuela.

References. Cameron and Fu (2006); Chen et al. 

(2006a, 2006b); Conran (1998); Judd (1998).

In Guaramacal, the family is easily recognized by its climbing habit, reticulate leaf venation, paired petiolar tendrils, unisexual flowers, and umbellate inflorescences.

Molecular, but not morphological, data generally place the 

Old and New World species of Smilax in separate clades (Cameron and Fu, 2006; Chen et al., 2006a, 2006b).

Smilax L.

Smilax L., Sp. Pl. 1028. 1753.

The characters of the genus are the same as those of the 

family.

References. Andreata (1997); Ferrufino-Acosta 

(2010); Gaskin and Berry (1998).

There is no current revision treating all of the neotropical 

species of Smilax. Proper identification of collections continues 

to be hampered by poorly prepared specimens (i.e., specimens 

lacking lower stems and/or population samples without representatives of both sexes), use of characters in treatments that are 

not reliable for delimiting species, and conflicting treatments of 

species depending upon political boundaries.

A KEY TO THE SPECIES OF SMILAX

1a. Stems terete, unarmed apically; leaf blades broadly ovate or ovate-lanceolate, nitid, margin and veins without prickles; 

tepals 3.5–7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. domingensis

1b. Stems angular or striate, armed or unarmed apically; leaf blades lanceolate or ovate-lanceolate, dull, margin and primary 

veins below with or without prickles; tepals 1.5–2.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. spinosa

Smilax domingensis Willd., Sp. Pl. 4(2): 783. 1806; Briceño, 

in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 

720–721, fig. 1 [as “2”]. 2010.

FIGURES 19, 26F

Smilax staminea Griseb., in Martius, Fl. Bras. 3(1): 11. 1842; Gaskin and 

Berry, in Berry et al., Fl. Venez. Guayana 9: 193, fig. 155. 2005.

Smilax floribunda Kunth, Enum. Pl. 5: 229. 1850, non Desv. ex Ham., 1825. 

Smilax kunthii Killip & C. V. Morton, Publ. Carnegie Inst. Wash. 461: 269. 

1936, nom. nov.; Dorr et al., Contr. U.S. Natl. Herb. 40: 59. 2000 [2001].

Smilax staminea f. obtusata Steyerm., Fieldiana, Bot. 28: 156. 1951.

Stems terete, glabrous, unarmed apically; terminal branches 

flexuous. Leaves broadly ovate or ovate-lanceolate, 7–11.5 × 

(3–)5–9.5 cm, apices acuminate, margins entire or slightly sinuate, bases acute to truncate, 3–5-veined, nitid; petioles 1.5–3 cm 

long. Inflorescences solitary. Peduncles 5–7 mm long; pedicels 

5–7(–10) mm long; tepals 3.5–7 mm long, tepals of staminate 

flowers slightly longer than those of pistillate ones. Berries maturing red, purple or black, ~1 cm in diameter.

Found in Mexico, Central America, the West Indies, and 

South America (Colombia, Venezuela, the Guianas, Ecuador, 
250 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 19. Smilacaceae, Smilax domingensis Willd. A. Habit. B. Inflorescence (pistillate). C. Flower (staminate). D. Flower (pistillate). 

E. Flower (pistillate), tepals removed and staminodes visible. F. Flower (pistillate), tepals and staminodes removed, stigmas visible. G. Leaf base 

with paired tendrils. H. Infructescence. (A, C, G, Dorr et al. 8916; B, D–F, Dorr et al. 9029; H, Dorr et al. 8900.)
NUMBER 100 • 251

Peru, Bolivia, and Brazil). In Venezuela, known from Amazonas, 

Aragua, Barinas, Bolívar, Distrito Federal, Mérida, Portuguesa, 

Sucre, Táchira, Trujillo, and Yaracuy (Map 310). Found in cloud 

forest on both slopes of Guaramacal; 1,800–2,500(–2,900) m.

Common Name. Bejuco chino.

Use. The stems are used to weave baskets (R. Caracas, 

pers. comm.).

Guaramacal collections of this species are consistently unarmed, but it is possible that these herbarium specimens are not 

complete and that prickles would be found at the base of the 

stems of plants in the flora.

Smilax spinosa Mill., Gard. Dict., ed. 8: [Smilax no. 8]. 1768.

Smilax scabriuscula Humb. & Bonpl. ex Willd., Sp. Pl. 4(2): 783. 1806.

Smilax mexicana Griseb. ex Kunth, Enum. Pl. 5: 167. 1850. 

Smilax scabriuscula var. fendleri A. DC., in Candolle and Candolle, Monogr. 

Phan. 1: 143. 1878.

Smilax sp. B; Dorr et al., Contr. U.S. Natl. Herb. 40: 59. 2000 [2001].

Smilax kunthii auct., non Killip & C. V. Morton; Dorr et al., Contr. U.S. 

Natl. Herb. 40: 59. 2000 [2001], pro parte (as to Steyermark & Rabe 

97375).

Stems slightly angled, striate, glabrous, armed with recurved 

prickles or unarmed apically; terminal branches zigzag. Leaves 

lanceolate or ovate-lanceolate, (7–)10–19 × 3–7(–9.5) cm, apices 

acute, margins entire or armed with prickles, bases rounded or 

subcordate, 5–7-veined, veins below often armed with recurved 

prickles, dull; petioles 0.7–1.3 cm long. Inflorescences solitary. 

Peduncles 2–3 mm long; pedicels ~1 mm long; tepals 1.5–2.5 

mm long, tepals of staminate flowers slightly longer than those 

of pistillate ones. Berries maturing purple to black, 5–7 mm in 

diameter.

Found in Mexico, Central America, and South America (Colombia, Venezuela, Trinidad and Tobago, the Guianas, Ecuador, 

Peru, and Bolivia). Widespread in Venezuela (Apure, Aragua, 

Bolívar, Carabobo, Cojedes, Distrito Federal, Falcón, Miranda, 

Monagas, Portuguesa, Sucre, Táchira, Trujillo, and Zulia; Map 

311). In the park, found on both slopes of Guaramacal; 1,900–

2,200(–2,900) m.

The name Smilax spinosa is applied to plants with relatively 

small tepals, peduncles shorter than petioles, and zigzag terminal 

branches. The type was collected in Mexico, and there is little 

agreement in the literature with respect to its range: Mexican and 

Central American floras generally omit South America, whereas 

Ferrufino-Acosta (2010) in her treatment of the West Indian species of Smilax includes northern South America and the northern 

Andes. In South America, especially in Venezuela and the Guianas, material that is very similar morphologically to S. spinosa

has been annotated as S. cumanensis Humb. & Bonpl. ex Willd., 

which Ferrufino-Acosta (2010), however, considers synonymous 

with the West Indian S. oblongata Sw. She separates S. spinosa

from S. oblongata on the relative lengths of peduncles and petioles; peduncles are shorter than petioles in the former species 

and equal to or sometimes longer than petioles in the latter. We 

believe this synonymy needs to be reexamined especially because 

the ranges of S. spinosa and S. oblongata overlap in the Cordillera de la Costa and in the Andes of Venezuela, and determinations of herbarium specimens by Ferrufino-Acosta (pers. obs.) 

suggest that S. oblongata also is found in the northern Andes.

MAP 310. Smilax domingensis occurrence in Venezuela. MAP 311. Smilax spinosa occurrence in Venezuela.
252 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Tofieldiaceae

L. J. DORR AND S. MIGUEL NIÑO

Perennial, rhizomatous herbs. Leaves simple, mostly basal, 

usually reduced upward on the stems and becoming bracts, linear, 

unifacial, entire or occasionally serrate, glabrous or inconspicuously tomentose, venation parallel, sometimes striate with a prominent central nerve and ribs. Inflorescences terminal, racemose or 

flowers solitary; usually bracteate. Flowers perfect, 3-merous, subtended by a calyculus (involucre) of (0)2 or 3(4) free to connate 

bracteoles; tepals 6, in 2 whorls, petaloid, free or basally connate. 

Stamens 6(9 or more), in 2 whorls; anthers basifixed or dorsifixed, 

2-thecate, introrse to latrorse, dehiscing by longitudinal slits; filaments free, flattened. Gynoecium of 3(4–6) carpels, nearly apocarpous or syncarpous below and apocarpous above, rarely totally 

syncarpous; ovary superior, 1-locular (apocarpous gynoecium) or 

3-locular (partially or totally syncarpous gynoecium); ovules few 

to many per locule; placentation parietal/marginal (apocarpous 

gynoecium) or axile (partially or totally syncarpous gynoecium); 

often with septal nectaries. Fruit a septicidal (or ventricidal) capsule or follicle-like. Seeds ellipsoid to fusiform with stipitate appendages, appendages sometimes reduced or absent.

A principally north temperate and circumboreal family of 

3 or 4 genera and ~30 species found in North America (Canada 

and USA), South America, Europe, and Asia. One genus, Harperocallis McDaniel, is found in the tropics and has an unusual 

disjunction occurring in the USA (Florida) and high elevations 

in South America.

References. Azuma and Tobe (2011); Remizowa 

and Sokoloff (2003); Remizowa et al. (2011); Tamura (1998); 

Tamura et al. (2004); Zomlefer (1997).

The genera of Tofieldiaceae have been placed in a number 

of different families, including Liliaceae s.l., Nartheciaceae, and 

Melanthiaceae, but molecular and morphological data now favor 

recognition of Tofieldiaceae as a distinct family. A morphological 

apomorphy is the presence of a calyculus, a structure not found 

in the other families mentioned. (The calyculus is sometimes secondarily lost in Tofieldia Huds.). Tofieldiaceae also can be distinguished from Nartheciaceae by fruit dehiscence (septicidal or 

ventricidal vs. loculicidal) and from Melanthiaceae by leaf orientation and shape (unifacial and equitant vs. bifacial and spiral).

Harperocallis McDaniel

Harperocallis McDaniel, J. Arnold Arbor. 49: 36. 1968.

Perennial herbs. Basal leaves distichous, equitant and unifacial, 

strongly ribbed, margins tomentose or rarely glabrous. Scape glabrous or rarely with eglandular hairs, 3–19 cauline leaves present, 

lowermost 1 or 2 well developed and others small and bract-like. 

Inflorescences a raceme, elongating in fruit. Flowers pedicellate, 

each pedicel subtended by a bract; each flower subtended by a calyculus of 3 free bracteoles. Tepals narrowly elliptic to narrowly 

obovate, persistent, connate basally, 5–9-ribbed. Stamens 6, free; 

anthers latrorse, basifixed; filaments free at base. Ovary ± oblong, 

2–3 mm tall, 3-locular; ovules numerous; styles fused almost to 

apex, apex usually truncate; stigmas 3, capitate. Capsules septicidal, 3–9-ribbed. Seeds fusiform, dark red with a white appendage 

at the chalazal end, decurrent along the seed body.

A genus of 10–12 species found in North America (USA) 

and South America. The South American species are found in the 

Andes (Colombia, Venezuela, Ecuador, Peru, and Bolivia) and 

the Guayana Highlands (Venezuela, Guyana, and Brazil). Eight 

or 9 species of Harperocallis occur in Venezuela.

References. Campbell and Dorr (2013); Cruden 

(1991).

Harperocallis is distinguished from other genera of Tofieldiaceae by its connate styles, pubescent leaf margins, and seed 

appendage that is decurrent along the seed body. The genus is 

known in recent literature as Isidrogalvia Ruiz & Pav., but that 

generic name is superfluous and illegitimate because it included 

the type species of Tofieldia when it was published. A nomenclatural proposal (Sokoloff et al., 2011) to conserve Isidrogalvia

with a conserved type was rejected (see Applequist, 2012: 1113).

Harperocallis robustior (Steyerm.) L. M. Campb. & Dorr, PhytoKeys 21: 47. 2013. Tofieldia sessiliflora var. robustior Steyerm., 

Fieldiana, Bot. 28: 157. 1951. Isidrogalvia robustior (Steyerm.) Cruden, Syst. Bot. 16: 279. 1991; Dorr et al., Contr. 

U.S. Natl. Herb. 40: 59. 2000 [2001]; Morillo, in Morillo et 

al., Bot. Ecol. Monocot. Páramos Venez. 2: 534–535. 2010.

FIGURE 20

Tofieldia sessiliflora auct., non Hook.; Ortega et al., BioLlania 5: 44. 1987.

Perennial herbs. Basal leaves sometimes falcate, 10–35 × 

0.5–0.85 cm, margins tomentose. Scape glabrous, 15–65 cm tall, 

with 0 or 1 well-developed cauline leaf and 7–17 bracts above 

MAP 312. Harperocallis robustior occurrence in Venezuela.
NUMBER 100 • 253

FIGURE 20. Tofieldiaceae, Harperocallis robustior (Steyerm.) L. M. Campb. & Dorr A. Habit. B. Leaf (detail of parallel veins and tomentose margins). C. Inflorescence (detail showing scape bracts and calyculus subtending flowers). D. Scape bracts (detail). E. Flower. F. Tepal. 

G. Ovary. H. Ovary (cross section showing axile placentation and ovules). I. Capsule. (A, I, Dorr et al. 4967; B–H, Stergios et al. 17327.)
254 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

the middle. Inflorescences generally elongated in fruit. Flowers 

few to several, greenish-yellow; pedicels 0.5–1 mm long (5–6 

mm long in fruit). Tepals 7–10 × 1–1.5 mm. Stamens slightly 

shorter than tepals. Styles separating at dehiscence. Capsules incompletely 9-ribbed.

Endemic to the Andes of Venezuela (Lara and Trujillo; Map 

312). In the park, found in the Páramo de Guaramacal, near 

the Laguna del Pumar, and on the summit of Cerro El Diablo; 

(1,950–)2,350–3,100 m.

Harperocallis sessiliflora (Hook.) L. M. Campb. & Dorr can 

be distinguished from H. robustior by the presence of eglandular 

hairs on the scape. Although the two species do not appear to be 

sympatric, the northernmost populations of the former, which is 

known from Colombia and Venezuela, approach the southernmost populations of the latter.

Xanthorrhoeaceae

L. J. DORR

Perennial herbs, rarely vines or nonwoody subshrubs, caespitose or rhizomatous. Roots fibrous and/or tuberous. Leaves mostly 

basal (rarely clustered at stem apex), distichous, unifacial, strictly 

bifacial or with an occluded region (i.e., bifacial both at the basal 

and distal ends of the leaf blade but in between just after diverging 

from the leaf sheath semiensiform and unifacial). Inflorescences 

variously branched panicles (or helicoid cymes), terminal, or flowers solitary; bracteate. Flowers actinomorphic or slightly zygomorphic, perfect, pedicellate, pedicels usually articulated. Perianth 

wholly or partially marcescent. Tepals 6, distinct or shortly connate into a tube, whorls subequal or inner slightly longer than 

the outer, variously colored. Stamens 6, free or slightly connate 

at base; filaments distinct, ornamented or not; anthers 2-loculed, 

basifixed, dorsifixed or centrifixed. Ovary superior or 1⁄2-inferior, 

3-locular (sometimes 3-locular below, 1-locular above); ovules 

few to many per locule; placentation axial or apical; style simple 

(rarely 3-parted apically), filiform; stigma terminal, punctiform to 

capitate. Septal nectaries present or not. Fruit a berry or capsule 

with complete or partial loculicidal dehiscence. Seeds ovoid; testa 

black, without appendages; endosperm copious.

A family of 35 genera and ~900 species found in South 

America, Europe, Africa, Madagascar, Asia, Australia, and the 

Pacific Islands. One genus is native to Venezuela, and several 

species-rich Old World genera are cultivated. At present, only 

1 species of these cultivated Old World genera is known to be 

naturalized in Venezuela.

References. Clifford (1998); Clifford et al. (1998); 

Smith and Van Wyk (1998); Wurdack and Dorr (2009); Zomlefer (1998).

Xanthorrhoeaceae comprises three well-defined subfamilies, 

including the Hemerocallidoideae to which the American genera 

belong and to which our family description applies. Eccremis 

Willd. ex Baker and Pasithea D. Don are the only Hemerocallidoideae native to South America, and despite their geographic 

proximity and the partial overlap of their respective ranges in 

southern Peru (at different elevations and apparently not sympatric), they are not closely related and clearly represent two separate introductions into the Americas. Eccremis, the genus found 

in this flora, is more closely related to Dianella Lam. ex Juss., 

which is native to Australia, Papua New Guinea, New Zealand, 

and other islands in the Pacific Ocean, than it is to its South 

American congener Pasithea (Wurdack and Dorr, 2009).

Hemerocallis L., an Asian genus of Hemerocallidoideae, is 

cultivated in Venezuelan parks and gardens but does not appear 

to escape. Species of subfamily Asphodeloideae, which are native 

to Africa, also are cultivated in Venezuela, and Aloe vera (L.) 

Burm. f., at least, is naturalized. This xerophytic species is found 

at elevations below 600 m and can be distinguished easily from 

Eccremis by its large rosette of succulent leaves and red flowers.

Eccremis Willd. ex Baker

Eccremis Willd. ex Baker, J. Linn. Soc., Bot. 15: 319. 1876.

Perennial herbs, to 2 m tall, rhizomatous or with short 

stems (not tuberous). Leaves clustered at the base, sheath flattened and flabellate, usually with an occluded region (appearing 

equitant), gradually reduced in size toward the apex; leaf blades 

linear-lanceolate, glabrous. Inflorescences lax corymbiform 

panicles; bracts leafy, conspicuous. Flowers showy, pendulous, 

pedicel articulated immediately below the flower; tepals prominently 5-nerved, to 10 mm long, blue (rarely white); stamens 6, 

minutely epitepalous, anthers centrifixed, filaments with a central swelling, minutely papillose, equal in length. Ovary superior; 

ovules numerous per locule. Stigma capitate. Capsule somewhat 

fleshy, loculicidal but only apically and not completely loculicidal, narrowly oblong, glabrous, pendulous; seeds ovoid.

A monotypic genus confined to South America, where it is 

found at high elevations in the Andes (Colombia, Venezuela, Ecuador, Peru, and Bolivia), the Cordillera de la Costa (Venezuela), 

and the Guayana Highlands (Venezuela and Brazil).

References. Ely and Luque Arias (2006); Schlittler 

(1940).

In the park, Eccremis can only be confused with Orthrosanthus (Iridaceae), from which it differs in having pendulous (vs. 

erect) flowers, 6 (vs. 3) stamens, glabrous (vs. pubescent) capsules, and ovoid (vs. flat) seeds.

Eccremis coarctata (Ruiz & Pav.) Baker, J. Linn. Soc., Bot. 15: 

320. 1876; Cruden, in Berry et al., Fl. Venez. Guayana 

6: 7–8, fig. 5. 2001; Morillo, in Morillo et al., Bot. Ecol. 

Monocot. Páramos Venez. 2: 506–508, fig. 1, foto 1. 2010. 

Anthericum coarctatum Ruiz & Pav., Fl. Peruv. 3: 67, t. 

299, fig. a. 1802.

FIGURE 26E

Dianella dubia Kunth, in H. B. K., Nov. Gen. Sp. [fol.] 1: 214. 1815 [1816]; 

ibid. [qu.] 1: 270. 1815 [1816]; ibid. [qu.] 7: t. 675. 1825.

Eccremis coarctata f. alba Steyerm., Fieldiana, Bot. 28: 153. 1951 

(“Excremis”).
NUMBER 100 • 255

The characters of this species are the same as those of the 

genus.

The distribution of this species is the same as that of the 

genus. In Venezuela, Eccremis coarctata is found only at high 

elevations (1,800–3,900 m) in the Andes (Lara, Mérida, Táchira, 

and Trujillo), the Sierra de Perijá (Zulia), the Cordillera de la 

Costa (Aragua, Distrito Federal, and Miranda), and the Venezuelan Guayana (Amazonas and Bolívar; Map 313). Páramo del 

Pumar; ~2,600 m.

The label on a specimen of Eccremis coarctata from Venezuela (Pittier 7578, US) suggested that the species was collected at 

200–400 m, an elevation well below those recorded for this species and in an unlikely locality (viz., Carabobo). Knuth (1928: 

199), however, indicated that the mislabeled specimen in question was in fact from Galipán (Distrito Federal) and collected at 

1,600–1,700 m.

Xyridaceae

L. J. DORR AND S. MIGUEL NIÑO

Perennial or rarely annual herbs, rosulate to caulescent, 

usually terrestrial, rarely aquatic (rooted never floating); caudex 

usually short; roots fibrous. Leaves alternate, simple, distichous 

or spiral; ligulate or not, bases broad; sheaths open, often equitant and keeled; blades laterally to dorsiventrally compressed, 

terete, angular or canaliculate. Inflorescences lateral or terminal, 

scapose; scapes 1 to few, arising from axils of scape sheaths or 

inner leaves, usually naked, each scape bearing at the apex 1(–2) 

imbricate-bracted spike or head or a panicle of spikes. Flowers 

perfect, 1 to many, subsessile to pedicellate in axils of chaffy, 

leathery or scarious bracts; perianth heterochlamydeous. Sepals 

(2) 3, unequal, anterior (inner) sepal rarely absent or weakly developed (Abolboda Bonpl. or Xyris L.), membranous and enveloping the corolla, usually caducous at anthesis (Abolboda and 

Xyris), the other 2 sepals subopposite, connivent to connate, 

chaffy to indurate, navicular, often keeled, clasping the mature 

capsule. Petals 3, equal or subequal, usually clawed, distinct or 

basally connate and tubular, salverform or porrect, yellow, blue, 

purple or white. Stamens 3(6), usually epipetalous, oppositipetalous; anthers basifixed, 2-thecate, 4-sporangiate, introrsely or 

laterally dehiscent, dehiscing longitudinally; staminodes (1–)3, 

clawed, distally 2-armed, plumose with moniliform or filiform 

hairs. Ovary superior, 1- or 3-locular, at least basally; ovules numerous, orthotropous or anatropous; placentation free-central/

basal or parietal (1-locular ovary); style 1, appendaged or not; 

stigmas 3-branched to U-shaped stigmas. Fruit a capsule, loculicidally dehiscent; seeds usually numerous, ellipsoid to globose, 

longitudinally striate to ridged.

An essentially pantropical family of 5 genera and ~400 species; 4 of the 5 genera, however, are restricted to South America, 

and the largest and most widely distributed genus Xyris also 

occurs in temperate regions of North America, Asia, and Australia. In the Americas, Xyridaceae are found in North America 

(Canada and USA), Mexico, Central America, the West Indies, 

and South America (Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina, and Uruguay). 

All 5 genera and ~100 species occur in Venezuela; approximately 

half of these species are endemic.

References. Campbell (2012); Idrobo (1954); Kral 

(1983, 1992, 1998); Rudall and Sajo (1999).

Molecular phylogenetic analyses of Xyridaceae do not resolve the family as monophyletic (Michelangeli et al., 2003; 

Davis et al., 2004), and if Xyridaceae s.l. is recognized (as is 

done here), there are clearly two major lineages. Morphology 

and palynology also tend to support this division (Kral, 1992, 

1998). The lineage comprising Xyris and Achlyphila Maguire & 

Wurdack (Xyridaceae s. str.) is marked by having usually distinct 

petals, distichous leaves, unappendaged styles, and pollen without spines or protuberances. The lineage comprising Abolboda, 

Aratitiyopea Steyerm. & P. E. Berry, and Orectanthe Maguire is 

united by having gamopetalous petals, polystichous leaves, styles 

with appendages, and pollen with spines or clavae. The latter 

lineage has been recognized at family rank as Abolbodaceae.

Xyris L.

Xyris L., Sp. Pl. 42. 1753.

Perennial or annual herbs; usually rosulate, solitary or caespitose. Leaves mostly equitant, distichous; ligulate; leaf blades 

usually laterally compressed, linear or sometimes terete or canaliculate. Scapes 1 to few per shoot, naked, terminating in 1 spike 

or head; bracts sometimes involucral, mostly chaffy. Flowers 

subsessile, solitary in bract axils. Sepals 3, 1 anterior (inner) 

sepal membranous, enveloping the corolla in bud, caducous at 

MAP 313. Eccremis coarctata occurrence in Venezuela.
256 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

anthesis, 2 lateral sepals navicular, free or connate with a variously margined keel. Petals 3, free (rarely connate), long-clawed, 

yellow. Stamens 3; filaments flattened and narrow; staminodes 

3 or 0, usually 2-branched, branches bearded. Ovary 1- or 

3-locular; placentation basal, central, or sometimes marginal, 

parietal or transitional; style elongate, 3-branched, each branch 

terminating in a U-shaped or semi-infundibuliform stigma; stylar 

appendages absent. Capsules membranous; seeds ovoid or fusiform, few to many.

Xyris with ~390 species is the most species-rich genus in the 

family, and it is found throughout the tropics and in temperate 

regions worldwide. Approximately 80 species occur in Venezuela, the majority confined to the Venezuelan Guayana.

Reference. Kral (1988).

Xyris subulata Ruiz & Pav., Fl. Peruv. 1: 46, t. 71, fig. b. 1798; 

Briceño and Campbell, in Morillo et al., Bot. Ecol. Monocot. Páramos Venez. 2: 728–730, fig. 2. 2010.

This species is composed of at least five varieties; the following is the most widely distributed variety and the only one 

known to occur in Venezuela.

Xyris subulata var. acutifolia Heimerl, Ann. K.K. Naturhist. 

Hofmus. 21: 63. 1906. Xyris acutifolia (Heimerl) Malme, 

Ark. Bot. 13(3): 40. 1913.

FIGURE 21

Perennial herbs, densely caespitose; stems contracted to 

slightly elongate. Leaves 10–14(–20) cm × (0.5–)1–2 mm, glabrous; sheaths entire, to 2–5 cm long, bases often shiny, pale to 

dark brown; ligules to 2 mm long or absent; leaf blades filiform to 

narrowly linear, flattened, margins smooth to papillate or scaberulous, apices narrowly acute. Scapes usually longer than the leaves, 

to 40 cm long, terete or oval in cross section. Spikes narrowly oblong to ovoid, 0.5–1 cm long. Petals mostly ovate, 3–5 mm long, 

yellow, apices rounded, coarsely erose. Staminodes 2-branched, 

branches flat, penicillate. Capsules ellipsoid, ~2.5–4 mm long.

Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, Guyana, Ecuador, and 

Peru). In Venezuela, recorded from the Andes (Lara, Mérida, 

Táchira, and Trujillo) and the Cordillera de la Costa (Anzoátegui and Sucre; Map 314). In the park, found in open, humid, and 

wet areas of the páramos of Guaramacal and Pumar, the Fila de 

Agua Fría, and other subpáramo areas; (2,350–)2,800–3,100 m.

Xyris subulata var. acutifolia is distinguished from the nominate variety by having entire (vs. ciliate) leaf sheath edges and 

contracted (vs. elongate) stems. Xyris subulata var. subulata is 

also Andean, but it does not occur in Venezuela and is known 

only from Colombia, Ecuador, and Peru.

Zingiberaceae

L. J. DORR AND B. STERGIOS

Perennial or rarely annual herbs; terrestrial (our species) or 

epiphytic; rhizomatous; aromatic. Aerial stems short, poorly developed or absent, unbranched; pseudostems (formed by overlapping sheaths) tall, generally conspicuous. Leaves distichous; 

sheaths usually open; ligule present or absent. Leaf blade entire; petiole present or absent; pulvinus usually absent (present 

in Zingiber Mill.). Inflorescences terminal on pseudostems or 

at the apex of short scapes arising from the rhizome and independent of pseudostems, lax or compact, few- to many-flowered 

monochasial cymes (cincinni); bracts of main axis enclosing or 

subtending flowers or cincinni. Flowers bisexual, epigynous, zygomorphic, heterochlamydeous. Calyx tubular, turbinate or urceolate, usually 3-lobed or 3-toothed, sometimes split down 1 

side. Petals 3, adnate basally to the androecium to form a narrow 

tube, the dorsal lobe larger than the other 2 lobes. Androecium 

highly modified, consisting of 1 fertile stamen (not petaloid), a 

2-lobed petaloid labellum, and sometimes 2 petaloid or toothlike, lateral staminodes, ± free or fused to the labellum; fertile stamen with a long or short filament; anther 2-thecate. Gynoecium 

3-carpellate; ovary inferior, 3-locular and placentation axile, 

or 1-locular and placentation parietal or basal; ovules many, ± 

anatropous. Style terminal, in a furrow of the filament and held 

between the 2 thecae; stigma funnelform, sometimes slightly 

2-lobed, slightly exserted from the thecae. Nectaries 2, epigynous. Fruit usually a capsule, loculicidally dehiscent from apex 

to base, 3-valved, irregularly dehiscent or indehiscent. Seeds few 

to many, ovoid to ellipsoid or subglobose, usually arillate.

A family of ~50 genera and 1,200–1,300 species found in 

the tropics and subtropics worldwide; centers of species diversity occur in East Asia and the Pacific Islands. In the neotropics, 

only the genus Renealmia L. f. is native. In Venezuela, species of 

the Asiatic genera Alpinia Roxb. and Hedychium J. Koenig are 

introduced and naturalized, and species of other paleotropical 

MAP 314. Xyris subulata var. acutifolia occurrence in Venezuela.
NUMBER 100 • 257

FIGURE 21. Xyridaceae, Xyris subulata var. acutifolia Heimerl. A. Habit. B. Scape (detail of surface). C. Leaf (detail of surface). D. Spike showing bracts and exserted apices of petals. E. Flower. F. Petal with adnate stamen. G. Plumose staminode. H. Apex of gynoecium. (A–C, Dorr et 

al. 8037; D–H, Dorr et al. 4950.)
258 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

genera such as Aframomum K. Schum., Curcuma L., Etlingera

Giseke, and Zingiber are cultivated in gardens.

References. Kress et al. (2002); Larsen et al. 

(1998); Rogers (1984).

The family is important as a source of commercial spices, including ginger (Zingiber officinale Roscoe), turmeric (Curcuma 

longa L.), and cardamom (Elettaria cardamom (L.) Maton and 

Amomum spp.).

KEY TO THE GENERA OF ZINGIBERACEAE

1a. Inflorescences terminal on pseudostems; bracts green; corolla white; lateral staminodes 3.5–4.5 cm long . . . . . Hedychium

1b. Inflorescences at base of plant on a short scape; bracts pink to bright red; corolla yellow to yellow-orange or orange; 

lateral staminodes 1–2(–3) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Renealmia

• Hedychium J. Koenig

Hedychium J. Koenig, in Retzius, Observ. Bot. 3: 61 [as “73”]. 1783.

Terrestrial (our species) or epiphytic herbs. Pseudostems 

well developed. Leaves sessile or petiolate; blades oblong or 

lanceolate; sheaths open; ligules conspicuous. Inflorescences 

terminal spikes; cincinni (1-) several-flowered; bracts broad and 

densely imbricate or narrow and exposing the inflorescence axis; 

bracteoles small, inconspicuous, usually tubular. Flowers white 

(our species), yellow or red. Calyx tube short, usually split on 1 

side. Corolla tube usually much longer than calyx and bracts; 

labellum large and showy, emarginate to 2-cleft; lateral staminodes 2, ± as long as the petal lobes, petaloid, and showy. Filament ± equal in length to labellum, usually much longer than the 

anther. Capsules 3-valved, globose or ellipsoid. Seeds numerous; 

aril lacerate.

Approximately 50–65 species native to Madagascar and 

Asia, with the greatest species diversity in tropical parts of the 

Himalayas and in Malaysia. At least 2 species have been introduced into Venezuela as ornamental plants, the following one 

now completely naturalized.

Reference. Wood et al. (2000).

• Hedychium coronarium J. Koenig, in Retzius, Observ. Bot. 3: 

61 [as “73”]. 1783.

FIGURE 22H,I

Erect herbs, 1–3 m tall, forming dense colonies. Leaf blades 

narrowly elliptic to oblong-lanceolate or lanceolate, (10–)20–60 

× 3–10 cm, glabrous above, finely pubescent below, sessile; ligules 1–4 cm long. Inflorescences dense, ellipsoid spikes, 5–20 × 

3–8 cm; cincinni (1) 2–6-flowered; bracts ovate to oblong, 4–5 × 

2–3 cm; bracteoles tubular, 30–35 mm long. Calyx tube 3–4 cm 

long, split on 1 side. Corolla tube very narrow, 10–12 cm long, 

white, but bright yellow in the center of the lobes, lobes linearreflexed; lateral staminodes 3.5–4.5 cm long, petaloid, white, 

fragrant. Capsules ovoid. Seeds black; aril red turning yellow.

Native to Asia. In the Americas, naturalized in North America (USA), Mexico, Central America, the West Indies, and South 

America (almost all country-level political units). Widespread in 

Venezuela (Amazonas, Aragua, Carabobo, Cojedes, Distrito Federal, Mérida, Miranda, Nueva Esparta, Portuguesa, Sucre, Trujillo, and Yaracuy; Map 315). Found near the lower boundaries 

of the park in Qda. Chanda on the north slope and near La Divisoria de la Concepción on the south slope; ~1,350 m.

Renealmia L. f.

Renealmia L. f., Suppl. Pl. 7, 79. 1781 [1782], nom. cons.

Terrestrial herbs. Leaves short-petiolate or sessile; sheaths 

open; ligules small; leaf blades ± narrowly elliptic. Inflorescence 

a raceme, terminating a separate, leafless shoot (our species) 

or sometimes terminal on a pseudostem; bracts herbaceous to 

membranous, striate, persistent, but then deciduous, enclosing 

a single flower (our species) or cincinni of 2–10 flowers; bracteoles herbaceous, tubular, and closed, or cyathiform and opening 

before anthesis (our species). Calyx tubular, turbinate or urceolate, 3-lobed, green, yellow, orange or red. Corolla tubular at 

base, exceeding the calyx, white or red, lobes imbricate in bud, 

1 upper lobe larger than 2 lateral lobes; labellum clawed, often 

pubescent, with 2 very small, toothlike lateral staminodes; limb 

(apical portion of labellum) composed of 2 rounded lateral lobes 

MAP 315. Hedychium coronarium occurrence in Venezuela.
NUMBER 100 • 259

FIGURE 22. Zingiberaceae. A–G. Renealmia thyrsoidea (Ruiz & Pav.) Poepp. & Endl. subsp. thyrsoidea. A. Habit. B. Inflorescence. C. Flower.

D. Flower (detail). E. Petal. F. Infructescence. G. Capsule. H, I. Hedychium coronarium J. Koenig. H. Flower. I. Inflorescence. (A, G, Dorr & 

Stergios 8851; B, Dorr & Yustiz 8547; C–F, Dorr et al. 8619; H, I, Angulo & Jeferson 15.)
260 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

and an entire or 2-lobed middle one. Stamen 1; filament shorter 

than labellum, ± equal to or shorter than anther. Ovary 3-locular, 

ellipsoid to globose; ovules few to many, 2-seriate. Capsules often 

black or red, 3-locular, ellipsoid to globose, ± fleshy, loculicidally 

dehiscent from base to apex, typically crowned by the persistent 

calyx. Seeds few to many; arillate.

A tropical American and African genus of ~100 species, of 

which ~75 are neotropical. In the Americas, found in Mexico, 

Central America, the West Indies, and South America (Colombia, 

Venezuela, Trinidad and Tobago, the Guianas, Ecuador, Peru, 

Bolivia, and Brazil). A dozen species are known from Venezuela.

References. Maas (1977); Särkinen et al. (2007).

The amphi-Atlantic distribution of Renealmia is remarkable 

as few angiosperm genera have significant numbers of species 

in both Africa and the neotropics. Molecular evidence indicates 

that Renealmia migrated from Africa to the Americas by the 

Miocene or Pliocene (15.8–2.7 mya; Särkinen et al., 2007).

Renealmia thyrsoidea (Ruiz & Pav.) Poepp. & Endl., Nov. Gen. 

Sp. Pl. 2: 26. 1838. Amomum thyrsoideum Ruiz & Pav., Fl. 

Peruv. 1: 2, t. 2. 1798.

Only the nominate subspecies has been collected in Venezuela and Guaramacal.

Renealmia thyrsoidea subsp. thyrsoidea

Maas, in Lasser, Fl. Venez. 11(2): 215–218, fig. 2. 1982.

FIGURE 22A–G

Erect herbs, 1–4(–5) m tall, forming colonies; rhizomes, 

crushed leaves, and floral parts all sweetly scented. Leaf blades 

narrowly elliptic, 35–55(–90) × 10–20 cm, bases cuneate, apices 

acuminate, midrib (when dry) darker than the blade, glabrous 

to sparsely covered with minute, bifid hairs above and below; 

petioles to 2.5–3 cm long; ligules 1–2 mm long. Inflorescences 

racemes, ~5–10 × ~3–5 cm, terminal on scapes arising from the 

rhizome and independent of the leafy stems; all parts of the inflorescence moderately to sparsely covered with small simple, 

forked or branched hairs; bracts ovate-obovate or obovatetriangular, 10–50 × 5–35 mm, pink to bright red; bracteoles 

cyathiform, 5–15(–20) mm long, open before anthesis. Calyx 

tubular, 10–25 mm long, yellow to red. Corolla tubular, 20–35 

mm long, yellow to yellowish-orange or orange, lateral lobes involute; lateral staminodes 1–2 mm long. Capsules 1.5–4 × 1–2.5 

cm. Seeds black; arils yellow to red.

Found in Central America (Costa Rica and Panama) and 

South America (Colombia, Venezuela, Trinidad and Tobago, Ecuador, Peru, Bolivia, and Brazil). Widespread in Venezuela (Amazonas, Anzoátegui, Aragua, Barinas, Bolívar, Cojedes, Distrito 

Federal, Falcón, Guárico, Lara, Mérida, Miranda, Portuguesa, 

Sucre, Táchira, Trujillo, Yaracuy, and Zulia; Map 316). In the 

park, found in various localities on both slopes of Guaramacal; 

1,600–2,100 m.

Common Name. Conópia.

Renealmia thyrsoidea subsp. chrysantha Maas, endemic 

to Guyana and Suriname, is distinguished from our subspecies 

mainly by the color of the bract and the calyx (golden-yellow 

vs. red), and also by its longer peduncles, pedicels, and calyces.

LIST OF NEW COMBINATIONS

Cyperaceae

Cyperus hortensis (Salzm. ex Steud.) Dorr, comb. nov.

Orchidaceae

Acronia archidiaconi (Ames) Carnevali & G. A. Romero, comb. 

nov.

Crocodeilanthe gelida (Lindl.) Carnevali & I. Ramírez, comb. nov.

Fernandezia schultesii (L. O. Williams) Carnevali & Dorr, comb. 

nov.

NOTE ADDED IN PROOF

Recently a narrower concept of Anathallis (Orchidaceae) 

than the one we adopted was proposed and the two species we 

recognized transferred to Stelis. While the author of this proposal 

argues that this creates a monophyletic Anathallis, the implications for Stelis are less clear and await further study. Karremans, 

A. P. 2014. Lankesteriana, a New Genus in the Pleurothallidinae 

(Orchidaceae). Lankesteriana 13: 219–332.

MAP 316. Renealmia thyrsoidea subsp. thyrsoidea occurrence in 

Venezuela.
NUMBER 100 • 261

FIGURE 23. A. Alstroemeriaceae. Bomarea amilcariana Stergios & Dorr. Inflorescence. B–J. Araceae. B. Anthurium amoenum Kunth & C. D. 

Bouché. Habit. C, D. A. nymphaeifolium K. Koch & C. D. Bouché. C. Leaf and inflorescence; note white spathe and red spadix. D. Leaf and 

inflorescence; note greenish-white spathe and yellow spadix. E. A. ramoncaracasii Stergios & Dorr. Leaf and inflorescence; note length of spadix 

relative to spathe. F. A. smaragdinum G. S. Bunting. Habit. G. A. ginesii Croat. Leaf base and inflorescence. H. Philodendron fraternum Schott. 

Inflorescence. I, J. Xanthosoma sagittifolium (L.) Schott. I. Inflorescence. J. Habit. (A–J, photographer B. Stergios).
262 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 24. A–C. Arecaceae. Wettinia praemorsa (Willd.) Wess. Boer. A. Habit and habitat (Qda. Segovia). B. Inflorescence. C. Infructescence. 

D–J. Bromeliaceae. D, E. Puya aristeguietae L. B. Sm. D. Habit and habitat (Páramo de Guaramacal). E. Inflorescence. F. Tillandsia fendleri

Griseb. Habit and habitat (slope forests of El Pumar). G. T. complanata Benth. Habit. H. P. venezuelana L. B. Sm. Habit and habitat (Páramo 

del Pumar). I. Mezobromelia capituligera (Griseb.) J. R. Grant. Inflorescence. J. Vriesea incurva (Griseb.) Read. Habit. (A, C–F, photographer 

L. J. Dorr; B, G, H, photographer S. M. Niño; I, J, photographer B. Stergios).
NUMBER 100 • 263

FIGURE 25. A. Cannaceae. Canna jaegeriana Urb. Inflorescence (detail). B. Cyclanthaceae. Sphaeradenia laucheana (Sander ex Mast.) Harling 

subsp. laucheana. Habit; note infructescence at base of plant (center left). C. Cyperaceae. Carex jamesonii Boott. Inflorescence. D. Eriocaulaceae. 

Paepalanthus pilosus (Kunth) Kunth. Habit. E. Heliconiaceae. Heliconia meridensis Klotzsch. Fruit. F. Orchidaceae. Elleanthus wageneri (Rchb. 

f.) Rchb. f. Inflorescence. G. Iridaceae. Sisyrinchium tinctorium Kunth. Flower and fruit. H–K. Orchidaceae. H. Anathallis sclerophylla (Lindl.) 

Pridgeon & M. W. Chase. Inflorescence. I. Epidendrum secundum Jacq. Inflorescence. J. Habenaria gollmeri Schltr. Inflorescence. K. Maxillaria 

triloris E. Morren. Flower. (A, C–E, G, H, photographer S. M. Niño; B, photographer L. C. Barnett; F, I–K, photographer B. Stergios).
264 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 26. A–C. Poaceae. A. Chusquea fendleri Munro. Branch. B. Andropogon bicornis L. Inflorescence. C. Cortaderia hapalotricha (Pilg.) 

Conert. Inflorescence. D. Orchidaceae. Prosthechea brachychila (Lindl.) W. E. Higgins. Inflorescence. E. Xanthorrhoeaceae. Eccremis coarctata

(Ruiz & Pav.) Baker. Inflorescence. F. Smilacaceae. Smilax domingensis Willd. Habit and inflorescences. (A, C, photographer S. M. Niño; B, E, 

F, photographer B. Stergios; D, photographer L. J. Dorr).
Acknowledgments

Numerous people and institutions contributed to the success of this project, and we 

are grateful to all of them for the many different ways in which they assisted us.

In Boconó, the staff of the Guaramacal National Park welcomed us and were generous with their time and knowledge. Amilcar Bencomo, superintendent, and Ramón 

Caracas and Luis Zambrano (deceased), park guards, were exceptionally helpful, and we 

could not have completed this project without their commitment and support.

In Guanare, the staff of the Herbario Universitario PORT and professors and students at the Universidad Nacional Experimental de los Llanos Occidentales “Ezequiel 

Zamora” (UNELLEZ) helped us collect, process, name, and ship specimens. We are especially indebted to Nidia L. Cuello A., Gerardo Aymard, and Angelina Licata D., who 

have been mainstays of the herbarium, an important national and international botanical 

resource. In nearby Mesa de Cavacas, Dorothy Stergios generously shared her home with 

visitors involved in this flora project.

In Washington, D.C., Rose Gulledge contributed more than any other individual 

to our success, and her contributions covered the entire spectrum of activities related to 

this flora project from database management to visitor logistics. Xochitl Munn provided 

bibliographic and editorial support. The staff of the U.S. National Herbarium managed 

the many specimen transactions and, with the help of volunteers, mounted the specimens 

that voucher this work. International collaboration would have been impossible without 

the assistance of two Smithsonian offices, the Office of Fellowships and Internships and 

the Office of International Relations. We are indebted to Roberta Rubinoff, Catherine 

Harris, Bruce (Will) Morrison (deceased), and Gordon Bullock, who handled fellowships 

and stipends, and Francine Berkowitz and Raymond Seefeldt, who helped with visas. In 

nearby Falls Church, Virginia, Larry Mendenhall provided inexpensive accommodation 

for the Latin American botanists involved in this project.

In Caracas, the Ministerio del Ambiente y de los Recursos Naturales (MARN) and 

the Instituto Nacional de Parques (INPARQUES) issued permits that allowed us to collect specimens in Guaramacal National Park. Also, the curators and staff of the Jardín 

Botánico de Caracas invariably extended a gracious welcome whenever we visited their 

herbarium.

In addition to the Herbario Universitario PORT and the U.S. National Herbarium 

(US), which have the most extensive collections of vascular plant specimens from Guaramacal National Park, we also consulted collections deposited in the Herbario Nacional de 

Venezuela (VEN), the Missouri Botanical Garden (MO), the New York Botanical Garden 
266 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

(NY), and the Orchid Herbarium of Oakes Ames (AMES). We 

are grateful to the curators of these herbaria for granting us access to Guaramacal and type material in their care.

We are grateful to the many botanists who examined our 

collections and provided expert determinations. They include 

Fernando Alzate Guarín (Medellín, Colombia), Jesús BotinaPapamija (Cali, Colombia), Eric Christenson (deceased), Lynn G. 

Clark (Ames, Iowa), Thomas B. Croat (St. Louis, Missouri), Gerrit Davidse (St. Louis, Missouri), Robert B. Faden (Washington, 

D.C.), Günter Gerlach (Munich, Germany), Jason R. Grant 

(Neuchâtel, Switzerland), Andrew J. Henderson (the Bronx, 

New York), Nancy Hensold (Chicago, Illinois), Bruce K. Holst 

(Sarasota, Florida), Emmet J. Judziewicz (Stevens Point, Wisconsin), W. John Kress (Washington, D.C.), Ximena Londoño 

(Cali, Colombia), Harry Luther (deceased), Mark Lyle (Jeetze, 

Germany), Ana Maria Molina (Buenos Aires, Argentina), Gerry 

Moore (Greensboro, North Carolina), Dan H. Nicolson (Washington, D.C.), Paul M. Peterson (Washington, D.C.), Ivón M. 

Ramírez (Mérida, Yucatán, Mexico), Héctor J. Rodríguez R. 

(Maracay, Venezuela), Gustavo A. Romero (Cambridge, Massachusetts), Zulma E. Rúgolo (Buenos Aires, Argentina), Robert J. 

Soreng (Washington, D.C.), Chelsea D. Specht (Berkeley, California), and Fred W. Stauffer (Geneva, Switzerland).

We are also grateful to the many librarians who helped us 

locate or verify publications. They include Pierre Boillat (Geneva, 

Switzerland), Robin Everley (Washington, D.C.), Peter Fraissinet 

(Ithaca, New York), Susan Fraser (the Bronx, New York), Sherry 

Vance (Boston, Massachusetts), and Judith Warnement (Cambridge, Massachusetts).

The line art was prepared by Cathy Pasquale and Alice 

R. Tangerini. The maps were designed and generated by Rose 

Gulledge with technical assistance from Ellen Farr and Sylvia 

Orli. All of the illustrations (line art and maps) were carefully 

checked and formatted for publication by Rose Gulledge.

We received financial support from the Andrew W. Mellon 

Foundation, Molson Breweries USA, Inc., the Consejo Venezolano de Investigaciones Científicas y Tecnológicas (CONICIT), 

UNELLEZ (Venezuela), and the Smithsonian Institution. 

UNELLEZ support was courtesy of the Secretaría de Investigación. Smithsonian support was courtesy of the Institution (the 

Atherton Seidell Endowment Fund and Scholarly Studies), the 

National Museum of Natural History (Biodiversity Surveys and 

Inventories Program, Research Opportunity Funds, and Small 

Grants), and the Department of Botany (Walcott Fund).

Fred W. Stauffer (Geneva, Switzerland) and Lisa M. Campbell (the Bronx, New York) reviewed the manuscript after it 

was submitted for publication. We greatly appreciate this timeconsuming and invaluable service and happily adopted their corrections and suggestions for improvement. The errors, of course, 

remain our own.
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Index of Scientific Names

Names of taxa recognized as occurring in the flora of Guaramacal are in bold type. All 

others are either synonyms or refer to taxa not present in the flora of Guaramacal. Page 

numbers in bold italic type indicate the start of the taxonomic account.

Aa, 109, 113, 160

hartwegii, 113

Abolboda, 255

Abolbodaceae, 255

Achlyphila, 255

Acianthera, 112, 114, 115

casapensis, 114, 115

chamensis, 114

prognatha, 114, 115

Acineta, 108, 115, 116

cryptodonta, 115, 116

erythroxantha, 115

sella-turcica, 115

Acraea, 101

multiflora, 101

Acronia, ii, 112, 116, 117, 118, 119, 120, 

121, 260

sect. Acronia, 116

sect. Macrophyllae-fasciculatae, 116

archidiaconi, ii, 117, 260

bivalvis, 117, 118

calamifolia, 116, 118

coriacardia, 117, 118, 119

phyllocardioides, 117, 119, 120

ruberrima, 116, 119

semiscabra, 117, 120

siphoglossa, 117, 120

subtilis, 118

tridentata, 120

sp. A, 117, 120, 121

Acrorchis, 163

Actinocephalus, 90

Actinopetala, 171

Ada, 123

Adamanthus, 124

Adeneleuterophora, 145

graminifolia, 145

Aechmea, 31, 32

paniculigera, 31

spectabilis, 31, 32

Aframomum, 258

Afrothismia, 44

Agapanthaceae, 8

Agrostis, 210, 211, 212, 213, 214, 220, 246, 

248

alba, 212, 213

indica, 248

meridensis, 211, 212

mertensii, 211, 212

perennans, 211, 212

pittieri, 211, 212

scabra, 211, 213

stolonifera, 213

subrepens, 212

venezuelana, 211, 213, 214

Aiphanes, 24, 25

lindeniana, 24, 25

stergiosii, 24

sp. A, 24

Alaticaulia, 171, 172

schlimii, 172

Alcantarea, 43

Alectoridia, 215

quartiniana, 215

Alliaceae, 8

Alloschemone, 21

Aloe, 254

vera, 254

Alpinia, 52, 256

spiralis, 52
282 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Alstroemeria, 5, 7

edulis, 7

Alstroemeriaceae, 4, 5, 6, 261

Altensteinia, 113

paleacea, 113

Amaryllidaceae, 4, 8

Amomum, 258, 260

thyrsoideum, 260

Amydrium, 19

Anacheilium, 188, 189

brachychilum, 189

lindenii, 189

Ananas, 31

Anathallis, 112, 121, 122, 260, 

263

acuminata, 121, 122

anderssonii, 121

dolichopus, 121

rubens, 121

sclerophylla, 121, 122, 263

Andropogon, 211, 214, 264

bicornis, 214, 264

 var. burchellii, 214

Andropogoninae, 214

Aneilema, 49

gracile, 49

Anthericum, 254

coarctatum, 254

Anthurium, 10, 11, 12, 13, 14, 

15, 16, 17, 18, 19, 261

sect. Calomystrium, 15, 16

sect. Dactylophyllium, 13

amoenum, 11, 261

 var. humile, 11

ascensus, 13

bernardii, 11, 12

bredemeyeri, 12

crassinervium, 11, 12, 13

eminens, 10, 13

 subsp. eminens, 13

 subsp. longispadix, 13

fernandezii, 12

gehrigeri, 11, 13

ginesii, 11, 14, 261

humboldtianum, 18

 subsp. viridispadix, 18

humile, 11

julianii, 11, 14, 15, 19

longegeniculatum, 11, 15

nubicola, 11, 14, 15

nymphaeifolium, 11, 15, 16, 

261

ramoncaracasii, 11, 16, 17, 

261

scandens, 10, 14, 16, 18

 subsp. pusillum, 18

 subsp. scandens, 16, 18

smaragdinum, 11, 15, 18, 

261

smithii, 12

subsagittatum, 18

wittianum, 13

Apatostelis, 196, 199, 200

chamaestelis, 199

hylophila, 200

Araceae, 1, 3, 9, 10, 17, 261

Aratitiyopea, 255

Arecaceae, 1, 3, 23, 24, 25, 53, 

262

Arecoideae, 24, 26, 27, 29, 30

Arthraxon, 211, 215

hispidus, 215

 var. hispidus, 215

quartinianus, 215

Arthrostylidium, 211, 215, 216, 

217, 232

geminatum, 232

pubescens, 216

venezuelae, 216, 217

Arum, 22

sagittifolium, 22

Arundinaria, 217

trianae, 217

Ascolepis, 61

Asphodeloideae, 254

Asplundia, 53, 54

moritziana, 53, 54

vagans, 53, 54

sp. A, 54

Aulonemia, 211, 217, 218, 233

hirtula, 233

trianae, 217

ximenae, 217, 218

Austroderia, 230

Aveneae, 229

Avetra, 86

Axonopus, 210, 218, 219, 242

compressus, 218, 219

Bactridinae, 24

Bambusoideae, 209, 211, 216, 

217, 224, 232

Baskervilla, 186

Benitzia, 44

suaveolens, 44

Blastocaulon, 90

Bomarea, 5, 6, 7, 8, 261

amilcariana, 5, 6, 7, 261

edulis, 5, 7

obovata, 7

ovata, 5, 7

pauciflora, 6

salicifolia, 5, 7, 8

truxillensis, 7

sp. B, 6

Borassus, 26

pinnatifrons, 26

Brachionidium, 113, 122, 123

sect. Yolanda, 122

steyermarkii, 123

tuberculatum, 123

Brachtia, 123, 124

glumacea, 124

sulphurea, 124

Brassia, 111, 123, 124

glumacea, 124

glumaceoides, 124

sulphurea, 124

Brassiopsis, 123, 124

glumacea, 124

Brevilongium, 182

Briegeria, 163

teretifolia, 163

Bromeliaceae, 1, 4, 30, 31, 262

Bromelioideae, 31, 32

Bromus, 210, 219, 220

catharticus, 219, 220

 var. catharticus, 219

 var. elatus, 220

 var. rupestris, 220

cebadilla, 220

Buccella, 171

Buesiella, 135

Burmannia, 44

Burmanniaceae, 3, 44, 86

Byrsella, 171

Caladieae, 22

Calamagrostis, 210, 211, 220, 

221

sect. Deyeuxia, 220

bogotensis, 220

chaseae, 221

effusa, 221

nuda, 220

pittieri, 221

planifolia, 220, 221

sp. A, 220, 221

Calawaya, 172, 174

meridensis, 174

Callisia, 49

gracilis, 49

Camaridium, 110, 124, 125, 126

bracteatum, 181

micranthum, 125

parviflorum, 125

purpureum, 125, 126

vestitum, 125

Campelia, 50

zanonia, 50

Canna, 45, 46, 263

jaegeriana, 45, 46, 263

leucocarpa, 45

paniculata, 45

Cannaceae, 3, 45, 263

Carex, 55, 56, 57, 58, 59, 60, 

62, 84, 263

acutata, 60

bonplandii, 57

hamata, 84

hermaphrodita, 62

jamesonii, 57, 58, 59, 263

longii, 57, 59

roraimensis, 57

tachirensis, 57, 60

tamana, 57, 60

Cariceae, 84

Carludovica, 53, 54

moritziana, 53, 54

Carludovicoideae, 52

Cartonematoideae, 46

Catoblastus, 30

praemorsus, 30

Catopsis, 39

fendleri, 39

Cenchrus, 211, 221, 222, 223, 

247

bambusiformis, 222

clandestinus, 222, 223

parviflorus, 247

peruvianus, 222, 223

Centrochloa, 218

Centropetalum, 158

Chaenanthe, 127

Chaetospora, 73

globosa, 73

Chaetotropis, 245, 246

elongata, 246

Chamaedorea, 24, 26, 27

linearis, 26

pinnatifrons, 26, 27

Chamaedoreeae, 26

Chaseopsis, 124

Chelyella, 124

Chillania, 64

Chimaerochloa, 230

Chloridoideae, 233, 248

Chusquea, 211, 216, 223, 

224, 225, 226, 227, 228, 

264

angustifolia, 224, 225, 228

aperta, 227

fendleri, 223, 224, 225, 264

glomerata, 226

magnifolia, 226

mollis, 224, 225, 226

multiramea, 224, 226

petiolata, 227

purdieana, 226, 227

serpens, 224, 227

silverstonei, 227

spectabilis, 224, 226, 227

spencei, 224, 228

tessellata, 224, 228

venezuelae, 216

Cinna, 210, 228, 229

poiformis, 229

Cladobium, 191, 192

Cleistes, 109, 126, 127

abdita, 127

costaricensis, 126

rosea, 126, 127

 f. pallida, 127

Cleistesiopsis, 126

Cocoseae, 24

Colanthelia, 217

Commelina, 46, 47, 50

hexandra, 47

obliqua, 47

robusta, 47

zanonia, 50

Commelinaceae, 4, 46, 48

Commelinoideae, 46

Commelinopsis, 46

Comparettia, 110, 127, 128

falcata, 127, 128

 var. falcata, 127, 128

 var. paulensis, 128

ottonis, 127, 128

stenochila, 128
NUMBER 100 • 283

Cordanthera, 204

andina, 204

Cornucopiae, 212

perennans, 212

Cortaderia, 210, 229, 230, 264

columbiana, 230

hapalotricha, 230, 264

roraimensis, 230

selloana, 230

Corymborkis, 108, 109, 128, 129

flava, 129

galipanensis, 129

Costaceae, 3, 50

Costus, 51, 52

guanaiensis, 51

 var. macrostrobilus, 51

 var. tarmicus, 51

macrostrobilus, 51

spiralis, 51, 52

 var. spiralis, 52

 var. villosus, 52

Cranichideae, 113, 129, 160, 

176, 186, 187, 191

Cranichidinae, 113, 129, 160, 

186, 187, 191

Cranichis, 109, 129, 130, 186, 

187

antioquiensis, 130

ciliata, 130

diphylla, 130

fertilis, 186

monophylla, 130

pycnantha, 187

stachyodes, 187

tenuis, 130

Crocodeilanthe, ii, 112, 121, 

131, 132, 133, 134, 196, 

260

elegans, 132

galeata, 132, 133

gelida, ii, 132, 133, 260

moritzii, 132, 133, 134

stergiosii, 131, 134

sp. A, 132, 134

Curculigo, 95, 96

capitulata, 95

Curcuma,158

longa, 158

Cyclanthaceae, 3, 52, 53, 263

Cyclanthoideae, 52

Cyclanthus, 52

Cymbidieae, 115, 124, 125, 

127, 135, 140, 159, 169, 

173, 177, 179, 180, 182, 

192, 203, 205, 206, 209

Cymbidium, 180

Cymbispatha, 50

Cymbocarpa, 44

Cymophyllus, 57, 84

Cyperaceae, ii, 4, 55, 58, 61, 

64, 67, 69, 71, 75, 85, 

101, 260, 263

Cyperus, ii, 56, 61, 62, 63, 64, 

260

densicaespitosus, 62

hermaphroditus, 62

hortensis, ii, 62, 63, 260

niger, 62, 63, 64

pumilus, 62, 63

Cyrtochilum, 111, 135, 136, 

137, 138, 139, 178

cimiciferum, 136

detortum, 136, 137

distans, 135, 136, 137

falcipetalum, 136, 137

flexuosum, 136

macranthum, 135

megalophium, 111, 135, 138

orgyale, 136, 138

ramosissimum, 136, 138, 139

undulatum, 138

zebrinum, 136, 138, 139

Dallwatsonia, 246

polygonata, 246

Danthonia, 210, 230, 231

hapalotricha, 230

secundiflora, 231

 subsp. charruana, 231

 subsp. matthei, 231

 subsp. secundiflora, 231

Danthonioideae, 229, 230

Dasyglossum, 135

Dendrobium, 122, 132, 180, 

208

acuminatum, 122

elegans, 132

pusillum, 208

Deyeuxia, 211, 220, 221, 229

bogotensis, 220

nuda, 220

planifolia, 221

poiformis, 229

Diadenium, 127

Dianella, 254

dubia, 254

Dichaea, 113, 139, 140, 141, 

142

sect. Dichaea, 140

sect. Dichaeastrum, 140

brachypoda, 140

camaridioides, 140

hystricina, 140, 141

latifolia, 140, 141

 var. longa, 141

longa, 141

morrisii, 141, 142

muricata, 141

robusta, 140, 141, 142

Dichanthelium, 209, 210, 231, 

232

acuminatum, 231, 232

 var. acuminatum, 231, 232

Dichorisandra, 46, 47, 48, 49

hexandra, 47, 48, 49

Didymogonyx, 211, 232, 233

geminatum, 232, 233

Dioscorea, 86, 87, 88

sect. Cylcadenium, 88

alata, 86

coriacea, 86, 87, 88

lehmannii, 86, 87, 88

lisae, 86, 88, 89

meridensis, 86, 88

moritziana, 87, 88

sp. A, 88

sp. B, 87

Dioscoreaceae, 3, 44, 84, 86, 89

Dioscoreales, 44, 86

Diptera, 164

Distichia, 101

Dracontium, 16, 19

pertusum, 19

scandens, 16

Dracula, 171

Dressleriella, 163

teretifolium, 163

Drymophila, 5

Duboisia, 177

reymondii, 177

Dupatya, 90

Eccremis, 98, 254, 255, 264

coarctata, 254, 255, 264

 f. alba, 254

Ecuadorella, 182

Egleria, 64

Eleocharis, 4, 56, 64, 65, 67

flavescens, 65

montana, 65, 66, 67

stenocarpa, 65, 67

Elettaria, 258

cardamom, 258

Elleanthus, 109, 142, 143, 144, 

145, 146, 147, 263

aurantiacus, 143, 145

columnaris, 143, 144, 146, 

147

confusus, 142, 144

flavescens, 143, 144, 145

furfuraceus, 143, 145

gracilis, 143, 145

graminifolius, 142, 145, 146

kermesinus, 144, 145

linifolius, 145

lupulinus, 143, 146

maculatus, 146

pusillus, 145

virgatus, 144

wageneri, 143, 146, 147, 263

Empusella, 195

endotrachys, 195

Encyclia, 188, 189, 199

subgen. Osmophytum, 188

sect. Euchile, 188

sect. Hormidium, 188

sect. Osmophytum, 188

brachychila, 189

ivonae, 199

lindenii, 189

Epidendreae, 114, 116, 121, 

123, 131, 147, 163, 167, 

168, 171, 177, 183, 185, 

188, 192, 194, 196, 206

Epidendroideae, 108, 114, 115, 

116, 121, 125, 127, 129, 

131, 140, 142, 147, 159, 

163, 164, 168, 169, 170, 

171, 173, 177, 179, 182, 

183, 185, 188, 190, 192, 

193, 196, 203, 205, 206, 

209

Epidendropsis, 147

Epidendrum, 110, 111, 113, 

125, 126, 147, 149, 150, 

151, 152, 153, 154, 155, 

156, 157, 158, 159, 163, 

188, 189, 263

subgen. Hormidium, 188

subgen. Osmophytum, 188

alpicola, 148, 149

anceps, 156

andinum, 148, 149, 150

attenuatum, 147, 150

aymardiorum, 152

bifarium, 149, 150

brachychilum, 189

breviracemum, 153

caesaris, 148, 150

cardioglossum, 158

cereiflorum, 148, 151

cernuum, 147, 155, 158

chioneoides, 149, 151

curtisii, 149, 151

dendrobii, 147, 152

elleanthoides, 153

fastigiatum, 156

frigidum, 152

frutex, 148, 152, 159

fruticetorum, 158

globiflorum, 150

guaramacalense, 148, 152, 

153

hartwegii, 189

ibaguense, 148, 153

jajense, 110, 147, 153, 154

klotzscheanum, 148, 154

lacustre, 148, 154

lansbergii, 151

leucochilum, 148, 154, 155

lindenii, 189

longiflorum, 154

macrostachyum, 157, 158

mojandae, 151, 158

obovatipetalum, 158

paniculatum, 156

 var. unguiculatum, 156

pichinchae, 158

pileatum, 152

pseudocernuum, 149, 155, 

158

ramosum, 157

repens, 148, 155, 156

restrepoanum, 150

secundum, 148, 156, 263

superpositum, 151

teretifolium, 163

unguiculatum, 148, 156, 157

urichianum, 149, 157
284 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Epidendrum (continued)

vareschii, 158

vestitum, 125

sp. A, 149, 157

sp. B, 149, 158

sp. C, 149, 158

Epipremnum, 19

Epithecia, 188

Eragrostis, 210, 233

tenuifolia, 233

Eriocaulaceae, 4, 90, 91, 263

Eriocaulon, 90

pilosum, 90

Eriochrysis, 210, 211, 234

cayennensis, 234

Erythrodes, 176

paleacea, 176

Etlingera, 258

Eucharis, 8, 9

amazonica, 9

grandiflora, 9

×grandiflora, 9

moorei, 9

sanderi, 9

Euchile, 188

Eulaema, 205

meriana, 205

Euterpeae, 29

Evelyna, 143, 144, 145, 146

aurantiaca, 143

columnaris, 144

flavescens, 144, 145

furfuracea, 145

gracilis, 145

lupulina, 146

wageneri, 146

Exalaria, 186, 187

parviflora, 187

Expedicula, 167, 168

apoda, 168 

Fernandezia, ii, 113, 158, 159, 

160, 260

crystallina, 159

pastii, 159

schultesii, ii, 159, 160, 260

Festuca, 210, 234, 235, 245

sect. Subulatae, 235

dinirica, 235

elviae, 235

guaramacalana, 235

Fimbristylis, 56, 68, 69

autumnalis, 68

 var. complanata, 68

complanata, 68, 69

Fissia, 171

Frondaria, 167

Fuertesiella, 191

Galeoglossum, 187

Geonoma, 24, 27, 28, 29

jussieuana, 27, 28

lehmannii, 27, 28

 subsp. corrugata, 27

 subsp. lehmannii, 27, 28

orbignyana, 27, 28

 subsp. hoffmanniana, 28

 subsp. orbignyana, 28

undata, 27, 28, 29

 subsp. undata, 28, 29

weberbaueri, 28

Geonomateae, 27

Glaziophyton, 217

Gomesa, 179

Gomphichis, 109, 113, 160, 

161

adnata, 160, 161

altissima, 160, 161

cladotricha, 161

costaricensis, 160, 161

gracilis, 160

steyermarkii, 160

traceyae, 161

viscosa, 160

Goodyera, 186

fertilis, 186

Goodyerinae, 176

Greigia, 31, 32

alborosea, 32

Guzmania, 31, 33, 34

mitis, 33

monostachia, 34

squarrosa, 33

Gymnosiphon, 44, 45

suaveolens, 44, 45

Gymnotrix, 222

bambusiformis, 222

Habenaria, 109, 130, 161, 162, 

163, 263

gollmeri, 162, 263

monorrhiza, 162, 163

parviflora, 130, 162

speciosa, 162

Haplothismia, 44

Harperocallis, 252, 253, 254

robustior, 252, 253, 254

sessiliflora, 254

Hedychium, 256, 258, 259

coronarium, 258, 259

Helcia, 205, 206

Heliconia, 92, 93, 94, 95, 263

bihai, 95

costanensis, 93

falcata, 93

hirsuta, 93, 94

latispatha, 95

meridensis, 93, 95, 263

schneeana, 93

stricta, 93, 95

Heliconiaceae, 3, 92, 94, 263

Helmia, 87

moritziana, 87

Hemerocallidoideae, 254

Hemerocallis, 254

Heteranthocidium, 179

abortivum, 179

Heteropsis, 21

Hexadesmia, 191, 192

Hexisea, 191, 192

Holochlamys, 21

Homalomena, 20

Homolepis, 210, 235, 236

glutinosa, 235, 236

Hormidium, 188

Humboltia, 201

oblonga, 201

Hypoxidaceae, 4, 95, 96, 97

Hypoxis, 95, 96, 97

decumbens, 96, 97

 var. major, 96

humilis, 96

Ichnanthus, 210, 236, 237

nemorosus, 236, 237

pallens, 236, 237

 var. majus, 237

tenuis, 236, 237

Irenea, 135

Iriarteeae, 30

Iridaceae, 4, 96, 98, 99, 100, 

254, 263

Isachne, 210, 238

ligulata, 238

rigens, 238

Isidrogalvia, 252

robustior, 252

Isotria, 126

Jacquiniella, 111, 163

teretifolia, 163

Jostia, 171

Juncaceae, 4, 101, 103

Juncus, 101, 102, 103, 104

bufonius, 102

densiflorus, 102

microcephalus, 102, 103, 104

Kalopternix, 147

Kobresia, 57, 84

Kyllinga, 61, 62, 63

hortensis, 62

pumila, 62, 63

viridiflora, 62, 63

Lachnocaulon, 90

Laeliinae, 147, 163, 188, 192

Lamprothyrsus, 229

Laricorchis, 180

Lasiacis, 210, 238, 239, 240

ligulata, 239

nigra, 239

procerrima, 239, 240

Lemnaceae, 9

Lemnoideae, 9

Lepanthes, 111, 164, 165, 166, 

167, 168, 207

acuminata, 165

barbae, 165

decipiens, 166

glicensteinii, 164

glochidea, 164, 165

lindleyana,165

millei, 165

scolex, 164, 165

stenophylla, 165

vareschii, 164, 165, 166

wageneri, 164, 165, 166

sp. A, 164, 166

sp. B, 164, 166, 167

Lepanthopsis, 111, 164, 167, 

168

apoda, 167

Leucohyle, 205, 206

Liliaceae, 252

Lindleyalis, 112, 168

glossopogon, 168

Lipocarpha, 61

Lockhartia, 113, 169

chocoensis, 169

Lolium, 235

Luerella, 171

Luzama, 171

Luzula, 101, 103, 104, 105

gigantea, 103, 104

racemosa, 105

Luzuriaga, 5

Luzuriagaceae, 5

Malaxideae, 170

Malaxis, 109, 169, 170

excavata, 170

licatae, 170

nidiae, 170

parthonii, 170

sp. A, 170

Maranta, 105, 106

arundinacea, 105

jacquinii, 105

lutea, 105

tonckat, 106

Marantaceae, 3, 105, 107

Mariscus, 62

hermaphroditus, 62

Marsippospermum, 101

Martinezia, 24, 26

lindeniana, 24

linearis, 26

Masdevallia, 112, 171, 172, 

173

subgen. Fissia, 171

civilis, 172

gerlachii, 171, 172

macroglossa, 172

schlimii, 171, 172

tubulosa, 171, 172

Maxillaria, 110, 120, 124, 125, 

172, 173, 174, 175, 180, 

181, 263

sect. Camaridium, 124

sect. Cucullatae, 124

sect. Ornithidium, 180

sect. Pseudomaxillaria, 124

sect. Reflexae, 180

sect. Siagonanthus, 180

aggregata, 181

calantha, 174

conferta, 125

luteoalba, 175

luteograndiflora, 175
NUMBER 100 • 285

macrura, 173, 174

 f. aurea, 174

meridensis, 173, 174

miniata, 181

nigrescens, 173, 174, 175

parviflora, 125

purpurea, 125

ruberrima, 181

semiscabra, 120

setigera, 175

triloris, 173, 175, 263

sp. A, 181

Maxillariinae, 125, 173, 180

Megema, 171

Melanthiaceae, 252

Melinis, 210, 240, 241

minutiflora, 240, 241

Mesospinidium, 123

Mezobromelia, 31, 34, 262

capituligera, 34, 262

Micrairoideae, 238

Microchilus, 109, 175, 176

paleaceus, 176

Millium, 218

compressum, 218

Minicolumna, 147

Monstera, 10, 19, 20, 21

adansonii, 19

 var. klotzschiana, 20

 var. laniata, 19

pertusa, 19

Monsteroideae, 10, 21

Moraea, 98

acorifolia, 98

Muscarella, 195

zephyrina, 195

Myoxanthus, 112, 176, 177

reymondii, 177

Myrosmodes, 113

Nartheciaceae, 44, 252

Nasonia, 158

Neobennettia, 169

Neodryas, 135

Neoescobaria, 205, 206

Neokoehleria, 127

Neourbania, 180

Neowilliamsia, 147

Neurolepis, 223, 225, 226, 227

aperta, 227

glomerata, 225

mollis, 225, 226

Nidularium, 32

alboroseum, 32

Niphantha, 133

gelida, 133

Ocampoa, 186

Ocellochloa, 210, 241

pulchella, 241

Octomeria, 184, 185

subgen. Pleurothallopsis, 184

Odontoglossum, 111, 135, 136, 

177, 178, 179, 182

subgen. Otoglossum, 182

brevifolium, 182

cimiciferum, 136

cinnamomeum, 178

constrictum, 178, 179

coronarium, 182

crocidipterum, 178

 subsp. dormanianum, 178

 f. dormanianum, 178

distans, 136

dormanianum, 178, 179

megalophium, 138

ramosissimum, 138

sanderianum, 178

schillerianum, 178, 179

zebrinum, 139

Oncidiinae, 124, 127, 135, 159, 

169, 177, 179, 182, 203, 

205, 206

Oncidium, 111, 135, 136, 137, 

138, 139, 178, 179, 180, 

182

subgen. Cyrtochilum, 135

sect. Cyrtochilum, 135

sect. Serpentia, 182

subsect. Serpentia, 182

abortivum, 179, 180

cimiciferum, 136

cinnamomeum, 178

crocidipterum, 178

subsp. dormanianum, 178

detortum, 136, 139

engelii, 139

falcipetalum, 137

orgyale, 138

zebrinum, 139

Oncodia, 123, 124

glumacea, 124

Ophiochloa, 218

Ophrys, 186

parviflora, 186

Oplismenus, 237

tenuis, 237

Orchidaceae, ii, 1, 4, 106, 108, 

260, 263, 264

Orchideae, 162

Orchidinae, 162

Orchidoideae, 108, 113, 129, 

160, 162, 176, 186, 187, 

191

Orchidotypus, 158, 159

schultesii, 159

Orchis, 162

monorrhiza, 162

Orectanthe, 255

Oreobolus, 56, 70, 71

venezuelensis, 70, 71

Oreodoxa, 29, 30

acuminata, 29

praemorsa, 30

Ornithidium, 110, 125, 126, 

173, 180, 181

confertum, 125, 126

miniatum, 180, 181

parviflorum, 125

ruberrimum, 181

serrulatum, 180

vagans, 181

vestitum, 125

Orthrosanthus, 98, 99, 100, 

254

acorifolius, 98, 99, 100

chimboracensis, 98

 var. acorifolius, 98

Otachyriinae, 247

Otoglossum, 110, 178, 182

coronarium, 182

Oxychloe, 101

Oxygyne, 44

Pachyphyllum, 158, 159

crystallinum, 159

hispidulum, 159

pastii, 159

schultesii, 159

Paepalanthoideae, 90

Paepalanthus, 90, 92, 263

subgen. Actinocephalus, 90

dendroides, 92

karstenii, 90, 92

 var. corei, 90

pilosus, 90, 91, 92, 263

Panarica, 188

Panicoideae, 214, 215, 218, 

222, 231, 232, 234, 235, 

236, 238, 240, 241, 242, 

246, 247

Panicum, 231, 235, 236, 237, 

238, 240, 241, 242, 246, 

247

subgen. Dichanthelium, 231

subgen. Phanopyrum, 247

sect. Lasiacis, 238

sect. Stolonifera, 241

acuminatum, 231

glutinosum, 235

nemorosum, 236

pallens, 237

 var. majus, 237

polygonatum, 247

procerrimum, 240

pulchellum, 241

rigens, 238

Pasithea, 254

Paspaleae, 247

Paspalum, 210, 218, 242, 243, 

244, 245

densum, 242, 243

distichum, 242, 243

inconstans, 242, 243

macrophyllum, 242, 243, 244

paniculatum, 242, 244

penicillatum, 242, 244, 245

prostratum, 244, 245

Patosia, 101

Pennisetum, 221, 222

bambusiforme, 222

clandestinum, 222, 223

longistylum, 222, 223

peruvianum, 223

Petalodon, 171

Pfitzeria, 127

Phaeosphaerion, 46

Phalaris, 215

hispida, 215

Phanopyrum, 247

gymnocarpon, 247

Philodendron, 10, 20, 21, 261

subgen. Meconostigma, 20

subgen. Philodendron, 20

subgen. Pteromischum, 20

fraternum, 20, 21, 261

rudgeanum, 21

tuerckheimii, 20, 21

Phyodina, 46, 49

gracilis, 49

Physothallis, 121

Physurus, 176

paleaceus, 176

Pilumna, 205, 206

laxa, 206

Pitcairnia, 31, 34, 35, 36, 37

brevicalycina, 35

caulescens, 35

feliciana, 35

hitchcockiana, 35

maidifolia, 35, 36

Pitcairnioideae, 34

Planotia, 227

aperta, 227

Platystele, 112, 182, 183

hypsitera, 183

pisifera, 183

Pleurothallidinae, 111, 114, 

116, 121, 123, 131, 164, 

167, 168, 171, 177, 183, 

185, 194, 196, 206

Pleurothallis, 112, 114, 115, 

116, 117, 118, 119, 120, 

121, 122, 131, 132, 133, 

134, 154, 167, 168, 177, 

183, 184, 185, 186, 194, 

195, 199, 207, 208

subgen. Acianthera, 114

subgen. Acuminatia, 121

subgen. Crocodeilanthe, 131

subgen. Specklinia, 194

sect. Lepanthopsis, 167

acuminata, 122

apoda, 167

archidiaconi, 117

biserrula, 168

bivalvis, 117

calamifolia, 118

casapensis, 114

chamaestelis, 199

chamensis, 114

coriacardia, 118

dependens, 207

dorrii, 184

elegans, 132, 133

endotrachys, 195

galeata, 132

gelida, 133

glossopogon, 168

grandiflora, 120
286 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Pleurothallis (continued)

lansbergii, 118

linguifera, 118

monocardia, 117

moritzii, 133

omoglossa, 117

phyllocardioides, 119

pisifera, 183

prognatha, 115

pusilla, 208

revoluta, 184

reymondii, 177

ruberrima, 119

sclerophylla, 122

semiscabra, 120

setigera, 195

siphoglossa, 120

stergiosii, 134

strobilifera, 184

subtilis, 118

tubulosa, 185, 186

vagans, 208

velaticaulis, 134

viridula, 185, 186

xanthochlora, 184

zephyrina, 195

Pleurothallopsis, 112, 184, 185, 

186

striata, 185

tubulosa, 185, 186

Poa, 210, 229, 233, 245

annua, 245

tenuifolia, 233

Poaceae, ii, 1, 4, 209, 220, 264

Pogonia, 126, 190

macrophylla, 190

rosea, 126

Pogonieae, 126

Pollardia, 188

Polypogon, 210, 245, 246

elongatus, 246

Ponthieva, 109, 186, 187

fertilis, 186, 187

parviflora, 186

Pooideae, 211, 219, 220, 229, 

234, 245, 246

Porphyrostachys, 113

Porroglossum, 171

Portillia, 171

Pothoideae, 10, 21

Pothos, 12

crassinervius, 12

Prescottia, 109, 187, 188

stachyodes, 187, 188

tubulosa, 187

Prestoea, 24, 29, 30

acuminata, 29

 var. acuminata, 29, 30

 var. dasystachys, 30

 var. montana, 30

Prioniaceae, 101

Prionium, 101

Prosthechea, 110, 154, 188, 

189, 264

subgen. Hormidium, 188

brachychila, 189, 264

lindenii, 189

Pseudencyclia, 188

Pseudocranichis, 187

Pseudocymbidium, 172

Pseudoludovia, 54

Pseudomaxillaria, 124, 125

parviflora, 125

vestita, 125

Psilochilus, 109, 190

macrophyllus, 190

physurifolius, 190

Psittacoglossum, 124

Pterichis, 109, 190, 191

diuris, 191

multiflora, 191

Puya, 31, 36, 37, 262

subgen. Puya, 36

subgen. Puyopsis, 36

aristeguietae, 36, 37, 262

maidifolia, 36

raimondii, 36

venezuelana, 36, 37, 262

Puyoideae, 36

Pycreus, 61, 63, 64

niger, 63

Queenslandiella, 63

Racinaea, 31, 37, 38, 39, 40

spiculosa, 38

 var. spiculosa, 38

tetrantha, 38, 39, 40

 var. caribaea, 39

 var. miniata, 39, 40

 var. scarlatina, 39

 var. tetrantha, 39

var. A, 39

Rajania, 86

Raycadenco, 158

Regalia, 171

Reichantha, 171

Reichenbachanthus, 191, 192

Reimarochloa, 242

Renealmia, 256, 258, 259, 260

thyrsoidea, 259, 260

 subsp. chrysantha, 260

 subsp. thyrsoidea, 259, 

260

Restrepiella, 185

tubulosa, 185

viridula, 185

Restrepiopsis, 185, 186

striata, 185

tubulosa, 185, 186

viridula, 185

Rhaphidophora, 19, 21

Rhipidocladum, 216, 232

sect. Didymogonyx, 232

geminatum, 232

Rhodohypoxis, 96

Rhodospatha, 10, 21

badilloi, 22

latifolia, 22

Rhoeo, 50

Rhynchospora, 56, 70, 72, 73, 

74, 75, 76, 77, 78, 79, 

80, 81

sect. Paniculatae, 81

aristata, 76

 var. immensa, 76

globosa, 72, 73

 var. globosa, 73

gollmeri, 72, 74

guaramacalensis, 73, 74, 75

hieronymi, 76

 subsp. hieronymi, 76

immensa, 72, 76

lechleri, 72, 76, 77

locuples, 72, 77

macrochaeta, 73, 76, 78, 80

polyphylla, 72, 78, 79, 81

rugosa, 72, 79

ruiziana, 72, 78, 80

tuerckheimii, 72, 79, 80

sp. A, 73, 81

Rodrigoa, 171

Rodriguezia, 128

stenochila, 128

Rondonanthus, 90

Rostkovia, 101

Rugoloa, 210, 246, 247

polygonata, 246, 247

Rusbyella, 135

Salmia, 54

laucheana, 54

Scaphyglottis, 110, 111, 125, 

163, 181, 191, 192

parviflora, 125

ruberrima, 181

summersii, 192

Scelochiloides, 127

Scelochilopsis, 127

Scelochilus, 127, 128

ottonis, 128

stenochilus, 128

Schlimmia, 108, 192, 193

alpina, 193

trifida, 193

Schoenoides, 70

Schoenoxiphium, 84

Schoenus, 78, 79

polyphyllus, 78

rugosus, 79

Sciaridae, 164

Scirpus, 65, 67, 68

complanatus, 68

flavescens, 65

montanus, 67

Scleria, 56, 81, 82, 83

distans, 82

tropicalis, 82, 83

Sepalosaccus, 124

Serapias, 129

flava, 129

Setaria, 210, 247, 248

geniculata, 247

parviflora, 247, 248

 var. parviflora, 247, 248

Setcreasea, 50

Siagonanthus, 180

Siederella, 135

Sigmatostalix, 179

Sisyrinchium, 98, 99, 100, 101, 

263

bogotense, 101

iridifolium, 100 

micranthum, 100

tinctorium, 99, 100, 101, 

263

Smilacaceae, 3, 249, 250, 264

Smilax, 249, 250, 251, 264

cumanensis, 251

domingensis, 249, 250, 251, 

264

floribunda, 249

kunthii, 249

mexicana, 251

oblongata, 251

scabriuscula, 251

 var. fendleri, 251

spinosa, 249, 251

staminea, 251

 f. obtusata, 251

sp. B, 251

Sobralia, 109, 193, 194

sect. Sobralia, 193

violacea, 193, 194

yauaperyensis, 193

Sobralieae, 142, 193

Spathiphylleae, 21

Spathiphyllum, 21

Specklinia, 112, 121, 133, 194, 

195

endotrachys, 194, 195

gelida, 133

zephyrina, 194, 195

Spectaculum, 171

Sphaeradenia, 53, 54, 55, 263

laucheana, 54, 55, 263

 subsp. irazuensis, 55

 subsp. laucheana, 54, 55, 

263

Sphagnum, 92

Sphenopholis, 229

Spilotantha, 171

Sporobolus, 210, 248, 249

indicus, 248, 249

jacquemontii, 249

Stanhopeinae, 115, 192

Stelis, 112, 121, 122, 131, 132, 

133, 195, 196, 197, 198, 

199, 200, 201, 202, 260

alba, 201

argentata, 197

ascendens, 196, 197, 198

atra, 196, 198

atroviolacea, 197, 198

bicornis, 199

biserrula, 196, 198, 199

chamaestelis, 196, 199

elegans, 132

galeata, 132

gelida, 133
NUMBER 100 • 287

hallii, 198

 var. atra, 198

humilis, 197, 199, 200

hylophila, 196, 200

jamesonii, 198

lindenii, 196, 200

moritzii, 133

neorubens, 121

nitens, 196, 201

oblonga, 197, 201, 202

picta, 132

purdiaei, 200

pusilla, 196, 201, 202

striolata, 196, 202

triseta, 198, 199

vulcani, 196, 202

Stellilabium, 203, 204

andinum, 204

pogonostalix, 204

Stenomeridaceae, 86

Stenomeris, 86

Stenoptera, 160

adnata, 160

Stenospermation, 21

Sterptoura, 171

Stigmatorthos, 127

Strelitzia, 92

Strelitziaceae, 92

Stromanthe, 105, 106, 107

jacquinii, 105, 106

tonckat, 105, 106, 107

Swallenochloa, 223, 224, 228

angustifolia, 224

tessellata, 228

Symphyglossum, 137, 177

distans, 137

Tacca, 86

Taccaceae, 44, 86

Tachinidae, 203

Takulumena, 147

Tamayorkis, 169

Tamus, 86

Telipogon, 109, 110, 111, 203, 

204

alticola, 204

andicola, 203

andinus, 204

astroglossus, 203, 204

bruchmuelleri, 204

latifolius, 203, 204

pogonostalix, 204

Tetragamestus, 191

Thecophyllum, 33

squarrosum, 33

Thismia, 44

Thismiaceae, 44

Thismieae, 44

Thrasyopsis, 242

Thurniaceae, 101

Tillandsia, 31, 34, 37, 38, 40, 

41, 42, 43, 262

subgen. Pseudocatopsis, 37, 38

sect. Pseudocatopsis, 37

aurantiaca, 39

 var. miniata, 39

biflora, 40, 41

capituligera, 34

caribaea, 39

compacta, 40, 41

 var. intermedia, 41

complanata, 40, 41, 42, 262

fendleri, 39, 40, 42, 262

 var. reducta, 42

incurva, 43

myriantha, 39, 40, 42, 43

rubra, 42

 var. reducta, 42

spiculosa, 38

tetrantha, 39

 var. caribaea, 39

 var. miniata, 39

Tillandsioideae, 33, 34, 38, 

40, 43 

Tiputinia, 44

Tofieldia, 252

sessiliflora, 252

 var. robustior, 252

Tofieldiaceae, 4, 252, 253

Tonina, 90

Tornelia, 19

laniata, 19

Tradescantia, 46, 48, 49, 50

gracilis, 49

zanonia, 48, 50

Tremarctos, 37, 42

ornatus, 37, 42

Trichocentrum, 110, 178, 205

pulchrum, 205

Trichopilia, 110, 205, 206

laxa, 206

sp. A, 206

Trichopodaceae, 44, 86

Trichopus, 86

Trichosalpinx, 111, 160, 164, 

167, 168, 195, 206, 207, 

208

apoda, 168

deceptrix, 207

dependens, 207

dunstervillei, 207, 208

pusilla, 160, 207, 208

zephyrina, 195

Trigonochilum, 135

distans, 137

Triotosiphon, 171

Triphoreae, 190

Tripogandra, 49

Trisetum, 229

Trisetella, 171

Tropidieae, 129

Typha, 234

Typhaceae, 234

Uncinia, 55, 57, 83, 84, 85

hamata, 84, 85

Vanilloideae, 108, 126

Vasqueziella, 115

Viracocha, 124

Vriesea, 31, 34, 43, 262

capituligera, 34

 var. lutea, 34

incurva, 43, 262

Vulpia, 235

Warrea, 209

cyanea, 209

Warreella, 108, 208, 209

cyanea, 209

Websteria, 64

Werauhia, 43

Wettinia, 24, 30, 262

praemorsa, 30, 262

Xanthorrhoeaceae, 4, 98, 254, 

264

Xanthosoma, 10, 22, 23, 261

pubescens, 23

sagittifolium, 22, 23, 261

Xyridaceae, 4, 255, 257

Xyris, 255, 256, 257

acutifolia, 256

subulata, 256, 257

 var. acutifolia, 256, 257

 var. subulata, 256

Yolanda, 122

Zahleria, 171

Zebrina, 50

Zingiber, 256, 258

officinale, 258

Zingiberaceae, 3, 256, 258, 

259

Zosterophyllanthos, 116

phyllocardioides, 119

Zygopetalinae, 140, 209

Index of Common Names

ave de paraíso, 92

bejuco chino, 251

bird’s nest anthurium, 12

capacho, 45

capin melao, 241

cat tail, 234

ciruelo, 52

conópia, 260

cucaracha, 47

espadilla, 99, 101

guaje, 23

guaje bravo, 23

guaje simangue, 23

guajesito, 13

junco, 68

kikuyo, 223

macanilla, 24

mapora, 30

moriche, 29

ñame de monte, 87

oso frontino, 37, 42

paja cebosa, 241

paja grama, 104

palmiche, 99

pampas grass, 230

pasto kikuyo, 223

piñuela, 37

platanillo, 92, 105

saladillo, 52

suelda con suelda, 47

virtud, 27
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