BOTANICAL MUSEUM LEAFLETS HARVARD UNIVERSITY PRINTED AND PUBLISHED AT THE BOTANICAL MUSEUM CAMBRIDGE, MASSACHUSETTS BOTANICAL MUSEUM LEAFLETS HARVARD UNIVERSITY VOLUME XVIII BOTANICAL MUSEUM CAMBRIDGE, MASSACHUSETTS 1957-1959 TABLE OF CONTENTS NuMBER 1 (June 7, 1957) The Identity of the Malpighiaceous Narcotics of South America By Ricuarp Evans SCHULTES. ....... 1 NuMBER 2 (June 26, 1957) The Effects of Certain Genes on the Outer Pistil- late Glume of Maize By WALTON C. GALINAP . Joe 4 «cu a « « BT Number 8 (January 27, 1958) Orchidaceae Neotropicales IV By Ricnarp Evans ScCHULTES . afi Studies in American Orchids III By Leste A. GARAY .... +. «s+. « » 108 A New Lepanthopsis from Venezuela By CuHarRLeEs SCHWEINFURTH ........ . 109 NumBeEr 4 (April 4, 1958) Plantae Austro-Americanae X By Ricuarp Evans SCHULTES. ...... .118 NuMBER 5 (November 20, 1958) How Absurd Can a Nomenclatural Proposal Be? By Lesuige A. Garay AND RicHarp Evans SOUDINGS . <4). Goel. se ew ee [v ] Studies in American Orchids IV By Lesuizg A. Garay ........ . . . 186 Two Additions to the Orchid Flora of Venezuela By CHarLes SCHWEINFURTH .. . . . . . . 219 A Little-Known Cultivated Plant from Northern South America By Ricuarp Evans Scuuures. . . . . . . . 299 Number 6 (January 16, 1959) New Considerations in an Old Genus: Datura By Arrnur S. Barchay ....... . . . 245 The Present Distribution of Narcotics and Stimu- lants Amongst the Indian Tribes of Colombia By Nesror Uscatrecur M. ..... . . . . 278 Plantae Colombianae X V By Ricuarp Evans ScHuutrEs. ..... . . 805 On the Validity of the Generic Name Cochleanthes Raf. By Ricnarp Evans Scuutres AND LesuiE A. GARAY ............2.2. 2.2. .821 NuMBER 7 (February 20, 1959) The Origin of Corn. I. Pod Corn, the Ancestral Form By Paun C. MaNGELsporr AND Rosertr G. REEVES. 2. 2...) B29 NuMBER 8 (March 10, 1959) The Origin of Corn. II. Teosinte, a Hybrid of Corn and Tripsacum By Roperr G. Reeves anp Patt C. MANGELSDORF . os It ~ [ vi ] Numser 9 (April 8, 1959) The Origin of Corn. III. Modern Races, the Prod- uct of Teosinte Introgression By Paut C. MANGELSDORF AND RosBert G. Wve faeces « & & «ahs (See BS BS Numser 10 (April 17, 1959) The Origin of Corn. IV. Place and Time of Origin By Paut C. ManGELSDORF AND Rospert G. REEVES. ...... ee ee ee ee ee ALB The Origin of Corn. V. A Critique of Current Theories By Ropert G. REEVES anpD Paut C. MANGELSDORF ........ +... + « 428 [ vii ] INDEX OF ILLUSTRATIONS PLATE Anthurium apaporanum PR. 17. Schultes. 2 2 . XIX Anthurium polyschistum RB. 2. Schultes & Idrobo LX Aristolochia georgica R. 7. Schultes . . . 2. XXVI Aristolochia xerophytica R. 7. Schultes . . XXVII Banisteriopsis Caapi (Spruce ex Griseb.) Morton II, [V, V, V1] Banisteriopsis inebrians Morton . . . . . . IDL, VI Banisteriopsis Rusbyana (Ndz.) Morton . . . . VII Bifrenaria minuta Garay... . . . OX Calliandra vaupesiana Cowan . . . . . . . XXIX Caularthron bicornutum (/Zo0ok.) Rafinesque X11,X 111 Caularthron bilamellatum (Pehb.f.) R. Ee. Schultes XIV, XV Conomorpha lithophyta R. 2. Schultes XXXI, XXXII, X XXIII Cyclanthus indivisus R. H. Schultes . . LVI, LIX Datura inoxia Miller 2...) CI Datura kymatocarpa Barclay . . . . . . LIL, LIV Datura meteloides DC. ew Dunal . . . . . . LL, Ll [ vill | Datura reburra Barclay... .. . . . LILI, LIV Datura vuleanicola Barclay... ... . LV, LVI Duckeella pauciflora Garay. . . . . . . . XXXIV Epidendrum superpositum Garay . . . . XXXVIII Hexadesmia Dunstervillei Garay . . . . XXXIX Inga involucrata Cowan . ....... XXVIII Inga longifolia Cowan . . . ...... XXVIII Jacqueshuberia amplifolia Cowan . . . . . XXIX Lepanthopsis vinacea C. Schweinf... . . . . XVII Lorostemon colombianum Maquire ..... XXX Makuna witch-doctor ........... =VIII Native tribes of Colombia who use stimulants and narcotics(Map) ........... LVII Notylia Norae Garay ........... XLII Octomeria deltoglossa Garay . . . . . . XXXVII Oncidium vagans C. Schweinf. . . . . . . . XLV Philodendron colombianum R. #7. Schultes . . . XX Platystele ornata Garay .........XXXVI Polyotidium Heubneri (Mansf.) Garay . . . XVI Rhytidanthera regalis R. H. Schultes. . . . . LXII Rodriguezia Teuscheri Garay... ..... XULI [ ix ] Solanum Topiro HBK. Adv, AVI, RLVIIL, XIX Stelis nexipous Garay... .. ..... XXXV Stenospermatium verecundum R. L2. Schultes XXI, XXII Telipogon Teuscheri Garay... .. . . XLII Tribes reported to use malpighiaceous narcotics (Map) I Urospatha antisylleptica R. WH. Schultes. . . . LXI Urospatha somnolenta R. 7. Schultes XXIII, XXIV, XXV Vargasiella venezuelana C. Schweinf. . . . . XLIV Zamia madida R. H. Schultes . . 2 2...) XVII Zea Mays L. Outer pistillate glumes . . IX, X, XI [x ] INDEX TO GENERA AND SPECIES AERANTHUS Lehmanni Rehb.f., 209,212 ALTERNANTHERA Lehmanii Hieron., 37,42,289 ambil, 289,291,293 ambird, 280,281,283 ambire, 280 AMMOPHILA arundinacea, 69 Andropogoneae, 57,67 ANTHOSIPHON Schltr., 214 ANTHURIUM apaporanum R, E.Schultes, 115,118,120 eminens Schott, 311 fontoides R. E.Schultes, 121 gracile (Rudge) Engl., 121 Harrisii (Grah.) Engl., 120 nanum FR. E.Schultes, 120 napaeum Engl., 120 polyschistum R.E.Schultes & Idrobo, 310,311,312 ApocyNacEAk, 15,39,42,113, 168,317 ARACEAE, 115,310,316 ARETHUSA sinensis Rolfe, 184 japonica A.Gray, 185 ARISTOLOCHIA cornuta Mast., 138 georgica R.E.Schultes, 125, 138,139,140 iquitensis O.C.Schmidt, 139 rumicifolia Mart. & Zucc.,142 sect. Exstipulatae, 138 sect. Pseudostipulosae, 139 sp., 18,42 xerophytica R. E. Schultes, 139,142,144 ARISTOLOCHIACEAR, 18,125 AVENA, 72 ayahuasca, 1,4—7,10,11,14-25, 30, 34,36,41,42, 290,292,293 ayawasca, 15,19,20 BACTRIS Sp., 291 BANISTERIA, 15,22,30,31 Caapi Spruce ex Griseb., 415 Jerruginea, 21 inebrians, 31 lutea Ruiz ex Griseb., 25 BANISTERIOPSIS, 7,16,20, 21,23,24,30,31,35,37,38,40, 41,42, 294 Caapi (Spruce ex Griseb.) Mor- ton, 4,5,7,8,10,11,14-17, 19-26,28, 30-32, 34,37-41 44,48,278 inebrians Morton, 12,21,22, 25,30, 35-37,41,45,46,278 289,290 longialata Ruis ex Ndz., 22, 30 metallicolor (4. Juss.) O°’ Don- nell & Lourteig, 24,25 quitensis (Ndz.) Morton, 19- 22,25,36,37,41,289 Rusbyanae (Ndz.) Morton, 22, 30,35,38,41,45,46,278, 289,290 spp., 230,292,296 barley mutants, 65 bejuco de oro, 36 be-td-ka, 242 BIFRENARIA, 208 grandis (Kriinzl. ) Garay, 208 minuta Garay, 206,207 picta (Schlir.) Schweinf., 208 sect. Lindleyella, 208 BLETIA, 182,163 florida R. Br., 185 Jormosana Hayata, 183 gebina Lindl., 185 hyacinthina (Sm.) R.Br., 185 kotoensis Hayata, 183 morrisonicola Hayata, 184 BLETILLA, 182,183 chinensis Schltr., 184 florida Rehb.f., 183,185 JSormosana (Hayata) Sehltr., 183 gebinae (Lindl.) Rehb.f., 183, 185 japonica (A.Gray) Schltr.,185 kotoensis (Hayata) Schltr., 183 morrisonicola (Hayata) Schltr. 184 ochracea Schltr., 184 scopulorum (W.W.Sm. ) Schltr., 184 sinensis (Rolfe) Schltr., 184 striata (Thunb.) Rehb.f., 185 sselschuanica Schltr., 185 yunnanensis Schltr., 185 var, Limprichtii Schltr. ,185 bo-po, 242 BOLLEA Wendlandiana Hort., 323 borrachero, 25,286,287 borrachera, 37,279,285 BROMUS, 72 Burseraceak, 156 caapi, 1,4-6,11,14—-16, 18,19, 21-25,30,39-42,293,295,298 caapi-pinima, 15,40,41 cagna, 16 CALANTHE gebinae (Lindl.) Lodd., 185 CALLIANDRA, 113,142,143 vaupesiana Cowan, 142,154 CAMARIDIUM, 208 equitans Schltr., 208 CAMPYLOCENTRUM Lehmannii (Rehb.f.) Schltr. , 212 Capanemieae, 104 capi, 6 CAPSICUM frutescens L., 230 CASSIA, 113,142 gigantifolia (Britt. & Killip) Cowan, 146 CAULARTHRON, 77 81,86,102 [ xii | bicornutum (Hook.) Raf., 82, 84, 86-90 bilamellatum (Rchb.f) R.E. Schultes, 86,88,89,90,92, 94,96-98, 100 umbellatum Raft, 80 CECROPIA peltata L., 292 spp., 292 CENCHRUS setigerus Vahl., 380 CENTRONIA laurifolia D.Don, 161 reticulata T7'riana, 160,161 vaupesana Wurdack, 160,161 chaco, 16 chagro panga, 30,35,36,46 cha-pe-nas, 291 CHAMAEDOREA sp., 291 CHAMAEFISTULA gigantifolia, Britt. & Killip, 146 chicha, 37 chimo, 280 CHONDRORHYNCHA, 321 aromatica (Rchb.f.) P.H. Allen, 323 calloglossa (Schltr.) P.H. Allen, 327 discolor (Lindl.) P.H. Allen, 324 Lipsicombiae Rolfe, 326 marginata (Rehb.f) P.H. Allen, 326 CLADONIA, 122 CLIBADIUM asperum (Auwbl.) DC., 230 coca, 229,278,280-288,29 1-299 COCHLEANTHES, 321,322 amazonica (Rchb.f. & Warsc.) R.E.Schultes & Garay, 322 aromatica (Rchb.f.) R.E. Schultes § Garay, 323 bidentata (Ichb,f. ex Hemsl.) R.E.Schultes §& Garay, 323 candida (Lindl.) R. E. Schultes & Garay, 323 digitata (Lem.) R. E.Schultes & Garay, 324 discolor (Lindl.) R. E. Schultes & Garay, 324 flabelliformis (Sw.) R.E. Schultes & Garay, 324 Jragrans Raf., 322,324 heteroclita (Poepp. & Endl.) R.E.Schultes & Garay, 325 ionoleuca (Ichb.f.) R.E. Schultes § Garay, 325 Klugii (C. Schweinf.) R.E. Schultes & Garay, 325 Lueddemanniana (Ichb.f.) R.E.Schultes & Garay, 326 marginata (Rehbf.) RE. Schultes § Garay, 326 trinitatis (4mes) R.E.Schultes & Garay, 326 Wailesiana ( Lindl.) R.E. Schultes § Garay, 326 cocona, 231,232,233,242 corn archaeological, 343-347 ,377-— 379,396-400,422—424 corn grass, 337,338, 352,430- 431 { xii | fossil pollen, 378,419—420, 423 modern races, 389-406 origin of, 329-440 current theories, 428-437 place, 413-423 time, 423-424 papyrescent, 339,340,352, 429-430 pod, 329-353 prehistoric, 343-347 reconstructing ancestor of, 349-351 CREPIS, 3865 sect. Ixeridopsis, 365 sect. Pyrimachos, 365 Cryptocentreae Garay, 213 CRYPTOCENTRUM, 212, Jamesonii Benth., 212 Lehmannii (Rehbf.) Garay, 209 pseudobulbosum Schweinf, 212 CUCURBITA Pepo, 282 CycaDACcKAR, 114 CYCLANTHACKAR, 305 CYCLANTHUS bipartitus Poir, 310 indivisus R. EF. Schultes, 305, 306,308,310 CYMBIDIUM flabelliforme (Sw.) Sw., 324 Alabellyfolium Sw. ex Griseb., 325 hyacinthinum Sm., 184 striatum (Thunb.) Sw., 184 DATURA, 245,285,286,289 arborea L., 22,279 candida ( Pers.) Safford, 279, 287 discolor Bernh., 259,260 dolichocarpa (Lagerh. ) Safford, 279,287 inoxia Miller, 245,247,252, 254,255 kymatocarpa Barclay, 256, 262,264 meteloides DC, ex Dunal, 245- 248 250,254,255 reburra Barclay, 258-260, 262,264 sanguinea Ruiz & Pav., 270, 271,279,287 sect. Stramonium, 260 sect. Dutra, 260 sp., 6,7,10,42 suaveolens H. & B, ex Willd., 25,279,285 vuleanicola Barclay, 260,266, 268,270,271 Wrighttt Regel, 246,247,254 dd-pee-ka-he, 168 de-twa, 242 DIACRIUM Benth., 77,79,80 amazonicum Schltr., 87,90 bicornutum (Hook.) Benth., 78,87,93,96 bidentatum (Lindl.) Hemsl., 102 bigibberosum (Rchb.f.) Hemsl. 92 bilamellatum (Rehb.f.) Hemsl. 93 var. Reichenbachianum Schltr., 93 bivalvatulum Schltr., 93,100 [ xiv ] indivisum (Bradf. ex Griseb. ) Broadway, 78,93,96,97 Ulmekei Krinzl., 102 venezuelanum Schltr., 93,100 djerabe, 291 DUCKEELLA Adolfii Campos Porto & Brade, 187 pauciflora Garay, 186-188 ELYONURUS tripsacoides, 67,72 EKPIDENDRUM, 77-81 bicornutum Hook., 73-80,86, 96,97 bidentatum (Lindl.) Hemsl., 102 bigibberosum Rehb.f., 92,96 bilamellatum Rchb.f. ,92,93,96 Boothianum Lindl., 102 flabelliforme Sw., 324 indivisum Bradf. ex Griseb., 92,96 obliquum Schiltr., 203 sect. Diacrium Lindl., 77,80 soratae Rchb. fi, 203 Steinbachii Ames, 203 stenopetalum Hook., 80 striatum (Thunb.) Thunb., 184 superpositum Garay, 203,204 tuberosum Lour., 184 ERIOCAULACEAE, 124 ERIOCAULON, 113 vaupesense Moldenke, 124,125 ERIANTHUS spp., 57 ERYTHRODES cylindrostachys Garay, 190, 191 quadrata Garay, 192 xystophylla (Rchb f.) Ames, 194 xystophylloides Garay, 193 zeuxinoides (Schiltr.) Ames, 193 ERYTHROXYLON Coca Lam., 229,278 novogranatense (Morris) Hieron., 278 EUCHLAENA, 57,379-381 EULOPHIA cochleata Knight ex Hooker, 324 cochlearis (Lindl.) Steudel, 324 Jloripondio 25 GOSSYPIUM, 368 grano turco, 419 guanguala, 282 guanguara, 282 guanto, 6 guay-ee-ga-m0-yoo-ke-ree, 39 guayaroche, 16 GUTTIFERAE, 113,158 GYAS humilis Salisb., 184 HACKELOCHLOA spp., 57 HAEMADICTYON, 15 amasonicum Bentham, 14,15, 40,42 hayawasca, 19 hayo, 280,281 [ xv ] HERRANIA nitida (Poepp.) R.E.Schultes, 230 HEXADESMIA Dunstervillei Garay, 203, 205,206 Jiminezii Schltr., 206 hi(d)-yati(d)yahe, 11 HOEHNEELLA Gehrtiana (Hoehne) Ruschi, 327 HUNTLEYA candida Hort. ex Lindl., 323 imbricata Hort. ex Rehb.f., 325 imbricata Pinel ex Lem., 324 marginata Hort. ex Rehb.f., $26 Huntleyinae, 321 HYBOCHILUS, 103 Huebneri Mansf., 1038-105, 108 tahi, 10 INGA, 113,142 inflata Ducke, 147 involucrata Cowan, 146,148 longifolia Cowan, 147,148 strigillosa, 150 IXERIS, 365 alpicola, 365 JACQUESHUBERIA, 113,142 amplifoliola Cowan, 150,154 quinquangulata, 151 JIMENSIA, 182,183 formosana (Hayata) Garay & R.E.Schultes, 183 kotoensis (Hayata) Garay & R.E.Schultes, 183 morrisonicola (Hayata) Garay & R.E.Schultes, 183 nervosa Rafi, 185 ochracea (Schltr.) Garay & R.E.Schultes, 184 scopulorum (W,W.Sm.) Garay & R.E.Schultes, 184 sinensis (Rolfe) Garay & R.E.Schultes, 184 striata (Thunb.) Garay & R.E.Schultes, 184 szetschuanica (Schiltr.) Garay & R.E.Schultes, 185 yunnanensis (Schltr.) Garay & R.E.Schultes, 185 kaapi, 11,18 kahée, 38 kahi, 24 kali, 11 katyja, 24 LACAENA grandis Krinzl., 208 LEGUMINOSAR, 142 LEPANTHOPSIS vinacea C.Schweinf., 109,110 LIMODORUM striatum Thunb., 184 LIPARIS elegantula Krédnsl., 202 neuroglossa Rcehb,f., 202 serpens Garay, 201,202 LOROSTEMON, 158 bombaciflorum Ducke, 158, 159 colombianum Maguire, 159, 162 Lorostemonoideae, 158 [ xvi ] lulo, 233-235 ma ma, 282 macuira, 279,280 MACAIREA albiflora Cogn., 165 arirambae Hub., 164,165 seabra Cogn., 165 Schultesii Wurdack, 164,165 theresiae Cogn., 165 MACROLOBIUM, 113,142 microcalyx Ducke var. minimum Cowan, 151 maicoma, 7,10 maikoa, 10 maize outer pistillate glumes, 57-76 normal glumes, 58-64 papyrescent glumes, 64-66 tunicate glumes, 66-68 vestigial glumes, 68-72 (See corn) maizillo, 395 MALAXIS tipuloides (Lindl.) O. Ktze., 201 var. exigua Garay, 201 MALOUETIA Tamaquarina, 39,42 MALPIGHIACEAR, 2,19,22,32,41 MANDEVILLA annulariifolia Woodson, 168, 169,170 Benthamii, 169 cuneifolia Woodson, 169 nerioides, 178 seabra, 176 stephanotidifolia Woodson, 176 Steyermarkii, 177 thevetioides Woodson, 177 manilla, 279 MANIHOT esculenta Crants., 229 utilissima, 283 MANISURIS, 72 spp., 67 MARSUPIARIA equitans Hoehne, 208 MASCAGNIA psilophylla (Juss.) Griseb. var. antifebrilis (Ruiz. & Pav.) Ndz., 19,20,41 MAXILLARIA, 208 equitans (Schiltr.) Garay, 208 heteroclita Poepp. & Endl.,325 Sophronitis (Rchb f.) Garay, 208 vandiformis Schweinf., 208 Maxillarieae, 212,213 Maydeae, 57,66,72,434 me-ne-ka-heé-ma, 38 MEeELASTOMACEAR, 113,160 METHYSTICODENDRON Amesianum R. E. Schulltes, 279,287 mets-kwat borrachera, 279 mihi, 11 Moronobeae, muémueli, 24 MyrsINAcKAE, 166 narcotics, 1—-56,273-304 nalema, 7,10,11,23,24,41 [ xvii ] nepe, 7,10,24,41 nepi, 7 NICOTIANA Tabacum L., 39,230,279 no, 280 noat, 280 NOTYLIA lilacina Arédnsl., 214 Norae Garay, 214,215 OcuNacrar, 317 oco-yagé, 30,35 ONCIDIUM vagans C.Schweinf., 223,224, 227 Warmingii Rchb f., 227 OCTOMERIA deltoglossa Garay, 199,200 rhodoglossa Schiltr., 199 ogfa, 36 ORNITHIDIUM Sophronitis Rehb.f., 208 Ornithocephalae, 213 pa-ree-ka, 294 pajuil, 39 Papperitzieae, 104 PARAPROTIUM, 113 amazonicum Cuatr., 156,157 vestitum Cuatr., 157 paricad, 278,294-296 PAULLINIA Yoco R. EF. Schultes, 279 PEGANUM Harmala, 22 pejt, 25 PENNISETUM ciliare (L.) Link, 380 PHILODENDRON colombianum R. E. Schultes, 121,126 PHREATIA, 212 PHYLLANTHUS spp., 220 PHYSOSIPHON, 195 PHYSOTHALLIS, 195 ptache, 279,280 pinde, 7,10,23,24,41 PIPTADENIA peregrina Benth., 278,295, 299,300 PLATYSTELE ornata Garay, 195,198 PLEIONE scopulorum W.W.Sm., 184 Pleurothallidinae, 195 POLYOTIDIUM Garay, 105 Huebneri (Mansf.) Garay, 105, 106 PRESTONIA amazonica ( Benth.) Macbride, 14,15,17,18,20,23-25,40, 42 Lindleyana Woodson, 179 vaupesana Woodson, 179 PROTIUM heptaphyllum, 297 pujana, 24 QUEKETTIA, 104 reé-ma, 38 RHYTIDANTHERA regalis I. F.Schultes, 317,318 roca, + [ xviii ] RODRIGUEZIA corydaloides Krinzl., 209 Teuscheri Garay, 209,210 sameruja, 6 Saundersieae, 213 SCELOCHILUS corydaloides (Kranzl.) Garay, 208 SEPALOSIPHON, 213 SIPHANTHERA capitata Gleason, 166 Hostmannii, 165,166 paraensis Huber, 166 pratensis Mg f., 166 ramosissima Cogn. ex Hoehne, 166 subtilis Pohl, 166 Tatei Gleason, 166 tenera Pohl, 166 vaupesana Wurdack, 165,166 snuff, 293-296,298-300 SOBRALIA candida (Poepp. & Endl.) Rehb f., 190 lancea Garay, 187,190 SOLANUM, 230 hyporhodium 4. Br. & Bouché, 232355 quitoense Lam., 233 Topiro HBK., 229-234,236, 238, 240-242, 244 SORGHUM, 65,66,74 papyrascens Stlapf., 65 sp.; 2383 STELIS, 195 nexipous Garay, 194,196 STENOSPERMATIUM verecundum R. FE. Schultes, 121,122, 128,130 Ulei, 122 sugii, 283 TABERNAEMONTANA coriacea Link ex R. & S., 180 rimulosa Woodson, 179,317 TACHIGALIA, 113,156 multijuga Benth., 153 polyphylla Poepp. & Endl., 153 Schultesiana Dwyer, 152,153 TELIPOGON semipictus Rchb.f., 218 Teuscheri Garay,214,216,218 Wallisii Rchb,f., 218 teosinte, 357-383,431-433 introgression, 389-406 TEPHROSIA toxicaria Pers., 230 TETRAPTERYS methystica FR. F.Schultes, 40, 41,278,298 sp., 33,41,42 THEOBROMA subincanum, 295 tobacco, 279-284,286,288,289, 29 1-296,298-300 ponga, 279,285 too-chée-ka-pee, 139 topiro, 233,234 TRIPSACUM, 57,69,72 australe, 370,402,403 dactyloides, 359,360,362, 369-371, 381 maizar, 370 [ xix | tupiru, 233,235 UROSPATHA antisylleptica R. EF. Schultes, 311,314-316 decipiens Schott, 123,316 Hostmannii, 316 sagittifolia (Rodach) Schott, 122 somnolenta RP. FE. Schultes, 123,124,132,134,136 Spruceana Scholt, 123 VARGASIELLA peruviana C. Schweinf. , 223 venezuelana C. Schweinf. , 219, 220 VIROLA calophylla Warburg, 278, 294,295 calophylloidea Markgraf, 278, 294,295 WARREA, 321 candida Lindl., 328 digitata Lem., 324 discolor Lindl., 321,324 marginata Rehb.f., 326 quadrata Lindl., 326 Wailesiana Lindl., 326 WARSCEWICZELLA, 321,322 amazonica Rehb.f. & Warsce., 322 aromatica (Rehb.f.) Rehb.f., 323 bidentata (Rchb.f. ex Hemsl.) calloglossa Schltr., 327 Schltr., 323 cochlearis (Lindl.) Rehb.f., 324 cochleata Barb.-Rodr., 325 digitata Barb.-Rodr., 324 discolor (Lindl.) Rehb.f., 324 Jlabelliformis (Sw.) Cogn. , 325 Gehrtiana Hoehne, 327 Gibeziae (N.E. Brown) Stein, 325 heteroclita (Poepp. & Endl. ) Hoehne, 325 ionoleuca (Rehb.f.) Schltr. 825 Lindenti Hort., 323 Lueddemanniacana Rehb.f. 826 marginata Rchb.f., 326 velata Rehb., 326 Wailesiana (Lindl. ) Rehb.f, ex Morren, 327 Wendlandi (Rehb.f.) Schltr. , 323 , ’ yageuco, 35 yagé, 24,25,30, 34-36 yageé cultivado, 36 yagé del monte, 35 yahé, 11,24 yaeé, 1,6,7,10,11,14, 16-25, 30, 33, 35-38, 41,42,44,46,278, 288-290, 292, 296,298,300 yajen, 14 ya-kee, 278,294 yako-borrachero, 25 yarumo, 292 yato, 278,294 yera, 291 yeras, 291 yoco, 279,288-300 yopo, 278,299 yuca, 229,283 [ xx ] yurupari, 25 ZAMIA madida R. F.Schultes, 114, 115,116 ZEA, 372,380,381 Mays L., 57 mexicana, 330,357,380,381, 389 Zygopetalinae, 321 ZYGOPETALUM, 321 amazonicum (Rehb.f. & Warsc.) Rehb.f., 323 aromaticum Rehb.f., 323 bidentatum Rchb.f. ex Hemsl., discolor (Lindl. ) Rehb.f., 324 flabelliforme (Sw.) Rehb.f., $25 Gibsiae N.E.Brown, 325 ionoleucum Rehb.f., 325 Klugii C.Schweinf., 326 Lindenii (Hort.) Rolfe, 323 Lueddemannianum (Rchb.f. ) Rehb.f., 326 makat H., 321 quadratum Pfitz, 326 rhombilabiumC. Schweinf. ,326 sect. Cochleare Rchb.f., 322 sect. Warscewiczella Rchb f., $22 trinitatis Ames, 326 323 Wailesianum (Lindl.) Rehb.f. candidum (Lindl. ) Rehb.f., 327 323 Wendlandi Rehb.f., 323 cochleare Lindl. , 321,322,324 cochleatum Paxt., 324 ya-to, 294 conchaceum Hoffmannsegg yato, 278 ex Rehb.f., 325 Issued June 3, 1959 Ee8 ERRATA Page 65, line 138 for Sholz read Scholz Page 105, line 19 for cornoso read carnoso Page 106 add Drawn by Leslie A. Garay Page 151, line 8 for basem read basim Page 123, line 6 for lanceoloto read lanceolato Page 172 reverse upper and lower legends Page 278 after Banisteriopsis inebrians add Banisteriopsis quitensis (Ndz. ) Morton, Colombia, Ecuador, Peru Page 283, line 7 fo utilissima read esculenta _ Page 310, line 1 for florium read florum Page 316, line 9 for Hostmanti read Hostmannii Page 317, line 18 fo specimine read speciminis = Page 317, line 26 for essensial read essential Page 323, line 22 for Gary read Garay = Page 324, line 1 for (Lehm.) read (Lem.) — Page 325, line 8 for Barb. Rod. read Barb.-Rodr. Page 375, line 14 for terminal read subterminal [ xxii | BOTANICAL MUSEUM LEAFLETS HARVARD UNIVERSITY VoL. 18, No. 1 CampBripGce, MassacnusetTts, JUNE 7, 1957 THE IDENTITY OF THE MALPIGHIACEOUS NARCOTICS OF SOUTH AMERICA BY Ricuarp Evans ScCHULTES This is all I have seen and learnt of caapi or aya-huasca. . . . Some travel- ler who may follow my steps with greater resources at his command, will, it is hoped, be able to bring away ma- terials adequate for the complete anal- ysis of this curious plant. Richard Spruce I Tue New World narcotics to which man has attributed the most extraordinarily bizarre powers of altering the state of his body and mind hid out successfully from pry- ing European eyes and were not discovered and identified botanically until about one hundred years ago. They remain, even to-day, the most poorly understood of the hallucinogens of the Americas and the narcotics whose identification is most baffling. Indeed, we may truthfully say that we stand merely on the threshold of our inves- tigations into the botany, ethnology, history, pharma- cology, chemistry and therapeutics of that complex of intoxicants known as ayahuasca, caapt or yaje. II The purpose of this paper is to summarize what Is known about the identity of the malpighiaceous narcot- [1] ics and to try to make some order from the rather chaotic state which at present obtains. A survey of the literature has been made, and the information from this source is evaluated in the light of field experience in the north- western part of the Amazon Valley which seems to be the centre of use of these narcotics. It is noteworthy that such a standard work as Hartwich (29) fails even to men- tion any intoxicant from the Malpighiaceae. This present study, however, leaves much to be de- sired. Since references to the malpighiaceous narcotics are widely scattered throughout botanical, anthropologi- cal, geographical and travel literature, I can make no claim to completeness of coverage. By far the greatest number of the references in regard to the sources of these narcotics are of limited or doubtful value; nevertheless it has been thought best to cite all that have come to light. I might say categorically, however, that much careful and sustained field work must be done before anything approaching a full understanding of the problem may be attained. Since this paper is concerned only with the identifica- tion of source-species, all discussion of the uses, effects, chemical constitution and ethnological importance of the narcotics themselves has been omitted. The line drawings were made possible by a grant from the American Academy of Arts and Sciences, and pub- lication of the illustrations has been done through a grant from the National Science Foundation. I express my appreciation to Miss Ruth Barton and Mr. Elmer W. Smith for the drawings reproduced herein. II] When, in 1851, the botanical explorer Richard Spruce penetrated the upper Rio Negro basin in Amazonian Brazil, a new and fertile epoch opened for natural science [2] PLATE I 3° _200 ace aaa geo = pr t re — 1 Guahibo 13 Bora 25 Sukuna | j 2 Zapare uXarihona 26. aiwane - s¥ngalere 15 uber aa ne ‘4 Mazan toSiukano 2Andoke »5 Correguahe wAbaku 2» Bare 6 Jivaro 1 Desano wHaniva 7 Colorado 9GYwanano 31 Jariana_, stayana rwekwano xfiyuka | *9JIngano wkakuna s3Ftoje _to-Jtona drasana dian 23 fanimuka Pit aNMira i _ TRIBES REPORTED TO USE | MALPIGHIACEOUS NARCOTICS (CAAPI, AYAHUASCA , YAJE) in the New World. Undoubtedly, one of Spruce’s great- est contributions to science was his discovery and ethno- botanical study of the curious caapt. He found the Tu- kanoan Indians of the Rio Uaupés and its affuents using caapt to induce for prophetic and divinatory purposes a narcosis characterized, amongst other strange effects, by frighteningly realistic colored visual hallucinations and a feeling of extreme and reckless bravery. This con- tribution included a precise identification, through flow- ering material, of a source of the narcotic, an hitherto undescribed species of a malpighiaceous liana. The liana, originally named Banisteria Caapi Spruce ex Griseb., is now correctly called Banisteriopsis Caapi (Spruce ex Griseb.) Morton. Although Spruce’s observations were written down in his wonted preciseness in 1852, they were not published until the posthumous account of his travels appeared in 1908 (90). In 1852, finding caapi cul- tivated in quantity on the Rio Uaupés, he noted: ‘‘There were about a dozen well-grown plants of caapi, twining up to the tree-tops along the margin of the roca [a cul- tivated plot] and several smaller ones. It was fortunately in flower and young fruit, and I saw, not without sur- prise, that it belonged to the order Malpighiaceae and the genus Banisteria, of which I made it out to be an undescribed species and therefore called it B. Caapi.”’ (90) Two years later, Spruce found caapi in use amongst the Guahibo Indians of the upper Orinoco in Colombia and Venezuela. Here the natives ‘‘not only drink an in- fusion, like those of the Uaupés, but also chew the dried stem, as some people do tobacco.’ Again, in 1857, whilst working in the Peruvian Andes, he encountered the Za- paro Indians using a narcotic known as ayahuasca. He stated that he ‘‘again saw caapi planted”’ in the north- eastern Peruvian Andes and that ‘‘it was the identical species of the Uaupés, but under a different name.’ (90) [ 4] The earliest published record of the use of any mal- pighiaceous narcotic of which I am aware dates from 1858, when Villavicencio (95) reported that the drug was em- ployed by the Zaparos, Angateros, Mazanes and other tribes of the upper Rio Napo in Amazonian Ecuador for sorcery, witchcraft, prophecy and divination. Although apparently no specimens were taken and no reference to a botanical determination was made in Villavicencio’s excellent and complete account, which included a report on self-intoxication, the common name ayahuasca was used, and the plant was described as a liana. Later work has definitely shown that ayahuasca in Peru and Ecuador belongs, in great part, at least, to the same genus as the Brazilian caapi. Indeed, as Spruce noted, although ‘‘of the plant itself?’ Villavicencio ‘‘could tell no more than that it was a liana or vine,’’ his ‘‘account of its proper- ties’’ coincided ‘‘wonderfully with what I had previously learnt in Brazil.’’ (90) In alist of plant names in the Tupi language of Brazil, the explorer von Martius (98,99) discussed caapi, stating that the Indians of the Rio Uaupés prepared from the fruits of the caapi plant an intoxicating drink, which they employed in their dance-ceremonies, and identify- ing it as Banisteriopsis Caapi. Since von Martius had never visited the Rio Uaupés, this report must be ac- cepted as an indirect one. He may have seen specimens of the caapi sent by Spruce to Europe and knew that Spruce had reported them as the source of the narcotic beverage. The statement that the fruits of the liana are used as the source of the intoxicants must be read with extreme reservation, even with doubt; for none of the reports of travellers and explorers who have seen the plant in use mention the fruits as the part of the plant employed. The few incidental references to caapi or ayahuasca [5] found in travel books have almost invariably neglected the fundamental question of the botanical identity of the source of the narcotic. Orton, who travelled widely in the Andes in the last half of the last century, reported (61) the narcotic use of ayahuasca amongst the Ziparos of the eastern slope of the Andes of Ecuador, but he failed to shed any light on the botanical source of the drug. Writing on his travels in South America, Crévaux reported (15) that, in southeastern Colombia, the Corre- guahes of the Rio Caqueta prepared an intoxicating bev- erage from the bark of a plant called yayé. He likewise noted, as had Spruce, the use by the Guahibos inhabit- ing the Rio Inirida of Colombia of an intoxicating drink prepared from a root called capi, apparently unaware that this capi and the yajé were probably botanically the same. Many years passed before it was known that yajé and ayahuasca referred to the same intoxicant. This report by Crévaux injected a new uncertainty into the picture. Was yajé from the same source as aya- huasca and caapi, or was it made from a different plant ? Although, as time went on, the belief that the three were at least generically identical became stronger, the prob- lem was not subjected to careful study until the late 1920s. In 1866, Simson reported (89) that the Piojes of Ecua- dor, like their neighbors, the Zdparos, ‘‘drink ayahuasca mixed with yajé, sameruja leaves and guanto wood, an indulgence which usually results in a broil between at least the partakers of the beverage.’’ He failed to iden- tify any of the ingredients botanically, but we may safely assume, I think, that gwanto refers to a tree-species of Datura. This report is, however, of great significance in being apparently the earliest to mention the mixture of anumber of elements in preparing the ayahuasea drink [ 6 | and the first intimation that yqj7é and ayahuasca might be different plants. In relation to Simson’s report that the Ziparos mix a number of plant materials to make the narcotic drink, we should point out Spruce’s definite statement that, so far as he could discover, the Banis- teriopsis was used alone by these Indians. Still a further complication arose when, in 1890, Ma- gelli, a missionary who had spent much time in the Pas- taza and Bobonaza areas of Amazonian Ecuador, reported (50) an intoxicating drink which the Jivaro Indians called natema. Concerning the botanical source of natema, Magelli wrote: ‘‘Natema is a red bark which, when boiled 24 hours, yields a decoction which deprives those who drink it of their senses for three days... .’’ Ma- gelli seems to have confused natema with another narcotic —maicoma—which he stated is ‘‘a little less toxic than natema.’’ It would appear from Magelli’s notes that he used the term natema to refer to a species of Datura and maicoma to refer to ayahuasca. As Reinburg (69) has suggested, Magelli undoubtedly erred and confused two plants whose uses were very similar. Later workers have established the fact that natema is actually referable to Banisteriopsis, whereas maicoma is, in reality, the much employed tree- Datura narcotic of the Jivaros. Tyler (92) reported, in 1894, the use and preparation of ayahuasca by the Zaparos of the Rio Napo area of Ecuador, stating merely that ‘‘the beverage is a decoc- tion of a certain species of liana. . .”’ and giving a de- scription of the effects of the drug. Writing of the Colorados of Ecuador in 1905, Rivet (73) referred to nepi (nepe) as a febrifuge, a violent emetic and an intoxicant and stated that its source was a “‘liana.”’ Later, in a glossary published in 1907 (74), he identified the Colorado nepe and the Cayapa pinde as Banisteriop- sis Caapi. We might here note that a more recent work ae EXPLANATION OF THE ILLUSTRATION Piate II. Banistertopsis Caapr (Spruce ex Grised. ) Morton. Drawing of a specimen of the type col- lection. 1, flowering branch, about one half nat- ural size. 2, flower, about two and one half times natural size. 3, fruit, somewhat larger than one half natural size. Drawn by Eimer W. Siri Prare fi BANISTERIOPSIS Caapi (Spruce ex Griseb,) Morton on these Indians by von Hagen (96) likewise refers the cultivated nepe to Banisteriopsis Caapi. It is again doubt- ful that any of these determinations were made on the basis of botanical specimens. In 1905, Rocha (77) published an account of his trip to the headwaters of the Rios Caquetaé and Putumayo in Colombia and reported that the Inga and Siona In- dians of the region, which to-day is included in the Comisaria del Putumayo, employed as a narcotic a “‘little bush” or ‘‘liana’’ called yajé. He stated that the natives would not allow white men to see the plant and that, for this reason, he did not know it from personal observa- tion. His account of its properties, however, coincided very closely with those described for ayahuasca, and it was widely assumed that the two were identical. In 1907, Rivet (75) wrote on the Jivaros of the Rios Pastaza and Bobonaza in eastern Ecuador. He indicated that they had a narcotic liana which they called natema and which he referred to Banisteriopsis Caapi. He clearly distinguished between the narcotics natema or yajé on the one hand and maicoma or maikoa (Datura sp.) on the other. Rivet’s identifications were apparently not based on botanical specimens. Later, Karsten (35) stated his belief that the ayahuasca of the Indians of the Rios Napo, Curaray, Bobonaza and Pastaza; the natema of the Jivaros; the pinde of the Cayapas; the nepe of the Colorados; and the caapi of the Rio Negro are all referable to one species: Banisteriop- sis Caapi. Ina letter to Reinburg (69), Karsten reiterated that he had no doubt that natema, ayahuasea, nepe and pinde are identical and may be referred to Banisteriopsis Caapi, even though he had available no identifications made on the basis of specimens, and that he could not offer a botanical determination of the ‘‘idhi’ (yajé?), an intoxicating liana which the Jivaros never used alone but [ 10 ] always mixed with natema. Furthermore, in 1926 (36) and again as recently as 1935 (87), he set forth his belief that ayahuasca and natema were the same and were refer- able to Banisteriopsis Caapi. Early in the present century, Koch-Griinberg, whose extensive travels (1903-1905) and investigations in the upper Rio Negro basin and in the Apaporis-Caqueta area are classic, stated that ‘‘kaapt’’ was prepared from a mal- pighiaceous shrub (39). I have been unable to find in our herbaria a specimen collected by Koch-Griinberg. Un- doubtedly basing his statement on Spruce’s earlier work in the same area, he referred the intoxicant to Banisteri- opsis Caapi. According to Koch-Griinberg (89), the Tukanos of the Vaupés distinguish two species of caapi; but for the second he ventured no botanical determina- tion. He also reported that the Karihonas (Hiandkoto- Umiua) of the headwaters of the Apaporis knew Banis- teriopsis Caapi, calling it yahe and hi(d)-yati(d)yahe. Since he did not penetrate to the sources of this river, he had to assume the ‘‘identification’’ without specimens. He stated, in a letter to Rivet (69) that, for the yayé of the Hiandkoto and the mihi of the Kubeos, he unfortunately did not have botanical knowledge and could not say whether it was Banisteriopsis Caapi or some other plant, but that he believed them both to be from this malpigh- iaceous species. During his studies amongst the Yekwanas of the upper Orinoco basin of southern Venezuela, Koch-Griinberg (40) found these people using two ‘‘species”’ of narcotic vines, one cultivated and one wild. Both were called kali. The stronger narcotic of the two was the cultivated kind. In his letter to Rivet (69) he stated his belief that these both represented Banisteriopsis Caapi. Hardenberg (28) and Whiffen (100), writing on the Colombian region lying between the Rios Caqueta and Ci ee EXPLANATION OF THE ILLUSTRATION Pirate III. Banisreriorsis ineprtans Morton. Draw- ing of the type specimen. Branch and fruit about one half natural size. Drawn by Etmer W. Siri [ 12 ] PuaTeE III BANISTERIOPSIS a tnebrians Morton SSRN . SSGGRAQAHyw SSQoon7yr sx SSS 7M) my S77! //) ) a mm! Putumayo, inhabited chiefly by Witoto and Bora In- dians, definitely reported the use of the narcotics in this area of the northwest Amazon, but neither was able to offer a botanical determination. Hardenberg stated merely that the liana was called ayahuasca or yajén; Whiffen, that it was known as caapi north of the Caqueta or ayahuasca south of this river. Both of these explorers thus intimated that these sundry names referred to one plant. In 1917, Safford (83), who had devoted much study to New World narcotic plants, referred both ayahuwasea and caapi to Banisteriopsis Caapi, after an evaluation of the literature on the subject. A suggestion that ayahuasca and yajé might be differ- ent plants, however, can be found in Reinburg’s study of the tribes inhabiting the region between the Rios Napo and Curaray in Pert. In 1921, he wrote (69) that the nar- cotic drink was an infusion of a few fragments of ayahu- asca, a liana the diameter of a man’s thumb, and leaves of yajé, “‘un petit arbuste, de 1m.50 de haut, 4 feuilles pétiolées (pétiole de 15 mm.) entiéres, ovales, longues de 20 cm., larges de 7 cm., réguliéres et terminées par une pointe de 2 cm.”’” On the basis of specimens collected, he held that ayahuasca, the source of which was always, according to his report, a wild, forest liana, and caapi were conspecific and represented Banisteriopsis Caapi. Still on the basis of specimens, he suggested that the yajé of the Curaray in Peru could, with reservation, be referred to the apocynaceous Haemadictyon (approach- ing, in some respects, Hl. amazonicum Benth.) or a re- lated genus. ' It was apparently Spruce (90) who first suggested that 'The generic concept Haemadictyon has been united with Prestonia, and the proper name of the plant to which Reinburg referred is now Prestonia amazonica (Benth.) Macbride (47). [ 14 | Prestonia amazonica might enter into the caapi picture. * He said that the eaapi-pinima (*‘ painted caapi’’) is ‘*an apocynaceous twiner of the genus Haemadictyon, of which I saw only young shoots, without any flowers. The leaves are of ashining green, painted with the strong, blood-red veins. It is possibly the same species . . . dis- tributed by Mr. Bentham under the name of Haemadic- tyon amazonicumn. sp. It may be the caapi-pinima which gives its nauseous taste to the caapi. . . and it is prob- ably poisonous... , but it isnot essential to the narcotic effect of the Banisteria, which (so far as I could make out) is used without any admixture by the Guahibos, Zaparos and other nations out of the Uaupés.”’ In 1922, ayahuasca or ayawasca was reported as a nar- cotie for the first time from Bolivia, from the Rio Beni, by White (101), thus greatly amplifying our knowledge of the range of the use of the intoxicant. White defi- nitely identified it as Banisteriopsis Caapi. Although he collected material for chemical analysis, it is not clear from his account whether or not his identification was based on an herbarium specimen. Much of White’s bo- tanical material is preserved in the Botanical Museum of Harvard University, but a search through the Eco- ? However, Spruce’s original field notebook, preserved at the Royal Botanical Gardens at Kew, shows a discrepancy, in this regard, with the published report. The field notes say: “9712, Banisteria Caapi Mss. From this is prepared an intoxicating drink known to all the natives on the Uaupés by the name of caapi. The lower part of the stem, which is the thickness of the thumb swollen at the joints, is the part used. This is beaten in a mortar with the addition of water and a small quantity of the slender roots of the Apocynac. (apparently a Haemadictyon) called caapi-pinima or painted caapi, from its lvs. being stained and veined with red . . . Query? May not the peculiar effects of the caapi be owing rather to the roots of the Haemadictyon (though in such small quantity) than to the stems of the Banisteria? The In- dians, however, consider the latter the prime agent, at the same time admitting that the former is an essential ingredient. The two plants rls are planted near all mallocas (villages)... [15 | nomic Herbarium of Oakes Ames at this institution fails to reveal a specimen of ayahuasca collected by White. In his report, White stated that the intoxicant was prepared either exclusively from the stems of ayahuasca, an immense liana with greenish white flowers, or else from ayahuasca stems boiled with the leaves of the chaco, a shrub with small, globose, red-yellow fruits. Leaves of plants locally called cagna and guayavoche may also be used with ayahuasca, but White could not ascertain whether these were other names for the plant known as chaco or whether they represented different species. It is unfortunate that so much uncertainty surrounds White’s report, but we may feel rather sure (from the common name ayahuasca, from his account of the intoxication and from recent plant collections in adjacent areas) that a species of Banisteriopsis is likewise the source of the nar- cotic in this Bolivian locality. In several reports, the botanist Rusby, who had accom- panied White on the Bolivian expedition, detailed the physiological effects of caapi and stated that the drug was derived from Banisteriopsis Caapi (80,81,82). Ina pharmacological report published in 1924, Seil and Putt (88) reported the isolation from Rusby’s material of a ‘‘fine powder”’ with at least three alkaloids (both phenolic and non-phenolic), but they offered no botanical deter- mination for the material which they had studied. In the same year, 1922, the Belgian botanist-explorer Claes, who had gone to the upper reaches of the Rio Caqueta, investigated the yaqjé of the Correguahe Indians of this area of southern Colombia (10). He learned that the yayé, hitherto usually described in the literature as ‘‘a small bush,’’ was an enormous forest liana. Claes argued—lI think quite correctly—that those who had de- scribed yajé as a small bush had seen young, cultivated individuals and not the vine in its wild state (9). [ 16 ] The intoxicating yajé of the Correguahe Indians was made by boiling pieces of the trunk itself together with the crushed bark of the trunk, but later the stem and leaves of a plant which the Indians did not permit him to examine were added to the mixture. This brew pro- duced, as Claes witnessed amongst the natives, very defi- nite narcotic symptoms. It is extremely unfortunate that Claes was unable to procure specimens of the leaves and stems which were used together with the liana, for we now know, from the studies of more recent investigators, that the narcotic yaje drink of this region of Colombia is sometimes of a more complex preparation than that of other parts of the Amazon; that very frequently a non- malpighiaceous ingredient may be employed. Claes did not offer a botanical determination of yajé (68). He mentioned that, according to De Wildeman, yajé ‘‘might be’? Prestonia amazonica. There is no speci- men of this species in the Rijksplantentuin in Brussels, and Claes himself states (68) that he did not obtain ma- terial for determining yajé, adding: ‘‘It thus is necessary to leave for others the task of collecting materials of the definitive classification of the plant.’” Michiels and Clin- quart (54), publishing their observations made during pharmacological experiments with Claes’ material, sug- gested that the stems with which they were working appeared to belong to Prestonia amazonica. The French pharmacologist, Rouhier, feeling (although apparently with no sound basis) that yajé and ayahuasca were one and the same thing, but that they might, physiologically, act differently, put forth the suggestion that ayahuwasca represents Banisteriopsis Caapi, whereas yqjé could rep- resent the unknown plant which Claes saw the Corre- guahe Indians add to the brew. A Colombian chemist, Fischer, was apparently the first to isolate a crystalline alkaloid from yajé (22). [17 ] Fischer admitted that botanical determination of yajé had not been made, but he stated that, to judge from anatomic and histologic details, it seemed to bea species of Aristolochia. In 1924, shortly after Fischer’s work was done, Rouhier (78) accepted this ‘‘identification,’’ stat- ing that his specimens ‘‘reminded one of a liana, proba- bly a species of the Aristolochiaceae.’’ In a subsequent work, however, Rouhier (79) attributed ayahuasca to Banisteriopsis Caapi but yajé to a ‘‘liana,’’ the identifica- tion of which was still under discussion. He pointed out that in its physiological action and its geographical range, yajyé was comparable to ayahuasca, and he dismissed its reference to Prestonia amazonica as ‘‘doubtful.’’ At about the same time, Barriga-Villalba (6), a Colombian chem- ist, and Albarracin (1), a Colombian pharmacologist, actively investigated yaqjé, but neither shed any further light whatsoever on the botanical identity of the drug. Albarracin described the source of his material as a ‘‘climbing shrub’’ and asserted that the natives did not cultivate yajé because it abounded in the forests. Reporting in 1926 on the caapi ceremony of the Tu- kano Indians of the Rio Tikié, a Brazilian affluent of the Uaupés, MacCreagh (49) described the intoxicant as ‘‘a thin, almost colorless liquid ... concocted from the leaves of a vine.’ He, apparently, failed to collect her- barium material of the vine. It is worthy of note that the Rio Tikié is the same area where Koch-Griinberg had made his observation on caapi. It is very near the locality where Spruce first met with the drug (90). Com- menting on ‘‘haapi,’’ which he had encountered in his travels in the same general region, the English explorer McGovern (53) failed to venture a botanical determina- tion, merely mentioning that the drug was made from **a root.” A rather inclusive survey of what had been accom- [ 18 | plished up to that date on the botanical identification and chemical constitution of this complex of malpighia- ceous narcotics was published in 1927 by two French pharmacologists, Perrot and Hamet (66,67). They pointed out that even more confusion reigned in the chemical field of investigation than in the botanical, prin- cipally because pharmacologists and chemists consistently disregarded taxonomic accuracy in identifying the sources of plant materials under study. Perrot and Hamet con- cluded that (a) yajé, ayahuasca and caapi refer to one species of plant: i.e., Banisteriopsis Caapi; and that (b) no apocynaceous plant is at all concerned in the problem of the source of the narcotic known under these three names. Notwithstanding the meritorious efforts of Per- rot and Hamet, their review brought little clarification into either the botanical or the chemical picture. In reply to the article by Perrot and Hamet, the Ger- man botanist Niedenzu (59) published several very in- teresting observations based on herbarium specimens preserved in the Berlin Herbarium. These specimens are, of course, no longer extant, but the notes are of special importance because of the authenticity of the determi- nations at the hand of an outstanding specialist in the Malpighiaceae. The collection Tessmann 5424 from Ya- rina Cocha on the Rio Ucayali in eastern Peru repre- sented, according to Niedenzu, a mixture: leaves of Mascagnia psilophylla (Juss.) Griseb. var. antifebrilis (Ruiz & Pav.) Ndz. and, in a paquet attached to the sheet, samaras of Banisteriopsis quitensis (Ndz.) Morton; on the label was annotated the vernacular name of haya- wasca or ayawasca and the observation ‘‘Stiicke der Liane gekocht, dann kalt genommen.’’ Niedenzu further called attention to three specimens of Banistertopsis quitensis. Eggers 15485, from eastern Ecuador, consisted of leaves, scanty flowers and plentiful fruit and had a note that, [ 19 ] from the stems, an intoxicating brew was prepared. Niedenzu stated that the characters of Mggers 15485 did not seem to correspond very closely with Banisteri- opsis quitensis as it was described. Tessmann 4974 and 5525, from the Rios Pastaza and Itaya, respectively, were referred without question to Banisteriopsis quitensis. Both were called ayawasca and were reported to be the source of a narcotic drink employed by witch-doctors. Tessmann identified Tessmann 5325 as Banisteriopsis Caapi, but Niedenzu pointed out that the collection was distinct from this species. In summary, Niedenzu stated that these wild and cultivated plants of Banisteriopsis from eastern Ecuador and Peru and the uses reported for them would seem to indicate that yageine, the alkaloid whose source was still in doubt, might well have come from similar species and that, on the basis of his studies, it would seem that three sources of the narcotic ‘‘aya- wasca’’ ought to be considered: Mascagnia psilophylla var. antifebrilis, Banisteriopsis quitensis and B. Caapi. Another attempt to put some order into the confusion resulting from field observations and fragmentary speci- mens occasionally gathered by ethnologists was made by Gagnepain, who, in 1930, published his botanical opin- ions relative to these narcotics (24). He pointed out (a) that, according to Reinburg, ayahuasca was ‘‘probably”’ Banisteriopsis Caapi, but that yayé could not be referable to this species; (b) that yayé seemed to approach Pres- tonia amazonica; (c) that fragments received as yajé by the chemist Rouhier in 1924 showed the plant to be an ‘““opposite-leaved vine’’; (d) that both Rivet and Rein- burg sent in material which seemed to represent the same malpighiaceous plant. Later, Gagnepain received, through Rouhier, a specimen from the Departamento de KI Valle in Colombia, where the plant was cultivated under the name of yajé.. The specimen, with leaves and [ 20 ] inflorescence attached, was determined as Banisteriopsis Caapi. This led Gagnepain to the rather extraordinary assertion that the yajé of Colombia was the same species as the caapi of Brazil, but different from the yayé of Ecua- dor. He could state with assurance only that the Ecua- dorian yajé represented a species of Banisteria ‘‘near ferruginea’”’ and that an exact clarification of the botan- ical sources of yayé was still far from realization. In 1929 and 1930, the Russian botanist Hammerman (26,27) published a rather complete survey of the prob- lem. His statements were founded on an evaluation of reports in the literature in the light of a study of material gathered in 1925-1926 by G. N. Varonof and S. V. Juzepezuk along the Rio Orteguaza, in the Colombian Comisaria del Caqueta. Pointing out that Zerda Bayén’s assertion that the Indians of the Caquetaé mixed four kinds of leaves to make a brew which they called yq7é, Hammer- man indicated that the material of Voronof and Juzepezuk gave a variety of results when chemically analyzed. He further stated that there seemed to be several species of Banisteriopsis involved, even though most of his material apparently was referable to Banisteriopsis quitensis. Any one of the species could be a source of the narcotic prep- arations of the Indians. He was, withal, careful to em- phasize that only Spruce had seen in flower an actual vine definitely known to be used to make the hallucinating drink. He intimated that the known variation in prep- aration, use and effect of the narcotics called caapi, yajé and ayahuasca might be due to differences in composition. In 1931, Morton (58) described a new species of Ban- isteriopsis from southern Colombia, naming it B.inebrians Morton because of its use by the Indians of the Com- isaria del Putumayo as an inebriant. Basing his state- ments on the meticulous field observations of the late Guillermo Klug, a German plant collector who worked [ 21 ] in eastern Peru and adjacent parts of Colombia, Morton indicated that at least three species are employed in this region: Banisteriopsis Caapi, B.inebrians and B.quiten- sis; and that Banisteria longialata Ruiz ex Ndz. and Banisteriopsis Rusbyana (Ndz.) Morton may enter cer- tain of the narcotic preparations as additional ingredients. Several reports on chemical studies of caap? published during the late 1920’s and the 1930's considered the bo- tanical identification of the intoxicant under investiga- tion. Reutter, reporting in 1927 (72) that he had isolated yageine and yagenine from the vegetal parts of yajé or ayahuasca, accepted Rouhier’s ‘‘identification’’ as Pres- tonia amazonica, stating that some tribes add to their yajé drink Banisteriopsis Caapi or ‘‘Datura arborea 1..”’ Lewin, a year later (44), wrote that ayahwasca was ‘‘a member of the Malpighiaceae.”’ In 1929, Keller and Gottauf (38) worked with material of ayahuasca from Bolivia and Peru, referring it to ‘‘a Banisteria,’’ (although they had no leaves nor flowers available) and isolated a harmine-like alkaloid. Costa and Faria (18,14) held that yajé, ayahuasca and caapt were the same narcotic and were derived from Bani- steriopsis Caapt. Three years later, Chen and Chen (8) summarized briefly the literature references and indicated their belief that caapi, yajé and ayahuasca represented Banisteriopsis Caapi, ‘‘a woody climber that attains a height of 8 to 4 metres and attaches itself to the trunks of large trees’’ or that is ‘‘grown in the North-western regions of South America.’’ Chen and Chen identified the alkaloids telepathine, yajeine and banisterine with harmine from Peganum Harmala, corroborating experi- mentally the earlier suggestions of Wolfe and Rumpf (104). The real importance of Chen and Chen’s paper, however, lies in the chemical study, apparently for the [ 22] first time, of vegetal material which seems to have been identified on the basis of botanical specimens. For the ‘‘twigs, leaves and roots of caapi’’ and the ‘‘decoction just as used by the Indians’ which were subjected to chemical analysis had been collected by the botanist Williams near Iquitos, Peru, and were determined as Banisteriopsis Caapi (4,102,103). For the most part, recent investigators who have de- voted critical attention to the study of the South Amer- ican malpighiaceous narcotics have concurred with Ham- merman, Gagnepain and Klug that several species of Banisteriopsis, if not of other genera, may be involved in some regions (12). Nevertheless, some contemporary writers tend to sim- plify the problem of identification, even though their ‘‘determinations’’ are seldom, if ever, based on speci- mens. While they are certainly not specialists, they are often quoted uncritically in technical works. Some, as in the case of Reko (70,71), have stated that ayahwasca, pinde, natema, caapi and yajé are all derived from Bani- steriopsis Caapit. In his dictionary of Amazonian plant names, Le Cointe (41) indicated a belief that ayahwasca and caapt were Banisteriopsis Caapi and that yaqjé corresponded to ‘‘another plant that enters into the composition of the caapi-drink as prepared by some tribes,’’ and he pointed out that some writers identify yajé as Prestonia amazon- ica. Von Hagen (96,97) considers both natema and caapi to be Banisteriopsis Caapi. In 1936, Pardal (62) referred caapi to Banisteriopsis Caapi and yajé to Prestonia amazonica. The following year, however, he stated (68) that caapi, yqjé and aya- huasca are Banisteriopsis Caapt. Maxwell (52), after an evaluation of the literature concerning the identity of caapi, concluded that the narcotic was a species of Ban- [ 23 ] isteriopsis, that ‘‘other vegetable ingredients are some- times included in the making of the beverage’ and that their identity is still in question. Lewin (42,45,46) identified natema, yagé, yahé, nepe, “*kahv’ and pinde as Banisteriopsis Caapi and enumerated the following Indian tribes as ‘‘addicted’’ to the use of this narcotic species: Guahibo, Tukano, Correguahe, Tama, Ziparo, Vaupé, Yekwana, Baré, Baniva, Manda- wake, Tariana, Siona, Jivaros, Colorados and Cayapas. Although Lewin held the principal ingredient to be Ban- isteriopsis Caapt, he admitted that it was sometimes used together with other plants. Amongst these plants, he named, as one possibility, Prestonia amazonica (43), al- though he strongly doubted that the name yajé referred exclusively to Prestonia amazonica. He further indicated that some Indians probably also add tobacco-water to the beverage. A most interesting note on ayahuaseca in Peru ap- peared in 1943 (94) and included a ‘‘recipe’’ for making the intoxicating beverage. Unfortunately, all of the plants employed were indicated with Indian names, with no hint as to their identity. The recipe commonly used in the vicinity of Iquitos has, as its principal ingredient, the liana or ‘‘death vine,’’ ayahuasca, which is undoubt- edly a Banisteriopsis. Into the decoction of this vine, the leaves of a species of muémueti are put; the name muém- ueti is said to refer to several plants, only one of which is used at atime. According to this account, the muémueti is responsible for the visual hallucinations. Alsoemployed in the decoction is an evil-tasting tuber called katija. To sweeten up the ayahuasca drink, leaves and seeds of a plant known as pujana are added. Herrera (31) listed the ayahuasca of Valle de Lares as Banisteriopsis metallicolor (A. Juss.) O’ Donnell & Lour- ‘planta t6xica.”” [ 24 ] teig and reported it as a Caller (7), in a consideration of the historical literature on the botanical sources of ayahuasca, concluded that the species used to prepare this narcotic drink is Banisteri- opsis Caapi and that B. quitensis is a synonym of B. Caapi. Other recent writers on Peru (18,97) attribute ayahuasea exclusively to Banisteriopsis Caapi. Padre Placido, who spent many years as a missionary in the Putumayo of Colombia, did not identify yajé botanically (17), but he stated that the Sionas often add to the narcotic drink of this name the pe or yako- borrachero—the floripondio of Mocoa, or, probably, Da- tura suaveolens H. & B. ex Willd. Sandeman (84) mentioned yajé casually, attributing it to Prestonia amazonica. Allen (2), in an excellent description of the yurupari ceremony of the Kubeo Indians of the Rio Vaupés in Colombia, indicated Banisteriopsis Caapi as the source of the narcotic beverage upon which this frenzied ritual is based. Taylor (91), following Allen, has attributed the malpighiaceous narcotics to Banisteriopsis Caapi. In arecent and authoritative work on plant alkaloids, Henry (30) identified the narcotics known as yajé, caapt and ayahuasca as Banisteriopsis Caapi, B. metallicolor or Banisteria lutea Ruiz ex Griseb. Manske and Holmes (51) attributed all three to Ban- isteriopsis Caapi. In 1946, Ducke (20) identified the fa- mous narcotic of the natives of the extreme northwest of the Amazon Valley, called caapi in the Brazilian part and yagé in the Colombian part of the region, as Bani- steriopsis Caapi. Hesse (82) attributed ayahuasca and yajé both to Banisteriopsis Caapi, and Moller (55) stated that caapi was exclusively referable to this species. Recently, Macbride (48), probably following Morton (58), pointed to Banisteriopsis Caapi, B. inebrians and B. quitensis as the principal sources of the alkaloid vari- [ 25 ] EXPLANATION OF THE ILLUSTRATION Pirate IV. Banistertopsis Caari (Spruce ex Griseb. ) Morton. Photograph of the plant from which the collection Schultes & Cabrera 18156a was made. Photograph by Hrrnanvo Garcia-Barrica [ 26 ] Prats TV EXPLANATION OF THE ILLUSTRATION Prare V. Cultivation of Banisteriopsis Caapi (Spruce ex Griseb. ) Morton (Schultes & Cabrera 17.209) by the Barasana Indians in the headwaters of the Rio Pira- parana, Comisaria del Vaupés, Colombia. Photograph by Ricuarn Evans Scuvvres PLATE V ously known as telepathine, yageine or banisterine. He stated, however, that it might also be found in ‘ta num- ber of related plants or forms** and quoted Morton in citing Banisteriopsis Rusbyana and Banisteria longialata as admixtures known by the names of oco-yagé and chagro panga. O’Connell and Lynn (60), working on material of Ban- istertopsis inebrians from the Putumayo of Colombia, found that the stems contain harmine and the leaves ‘‘an alkaloid which was partly identified as harmine.’’ The work of O’Connell and Lynn is noteworthy as represent- ing apparently the second chemical investigation carried out on material of Banisteriopsis identified through her- barium material. Mors and Zaltzman published a most interesting con- tribution in 1954 (56) in which they indicated their opin- ion, based on chemical examination, that yageine was different from harmine. On the basis of a brief review of the literature, they concluded that eaapi and ayahuasea were referable to Banisteriopsis Caapi but that yagé was not the same narcotic. Most recently, Fabre (21) published an historical re- view of caapi, ayahuasca and yajé, indicating his belief that only one species Banisteriopsis. Caapi--is_ the source of these narcotics, though citing Spruce that, amongst certain tribes, another plant may oftentimes be added in the preparation of the narcotic drink. IV Although Morton (57) has convincingly shown why the generic epithet Banisteria is not available for the group of malpighiaceous plants which should be called Banisteriopsis, in accordance with the lucid arguments of Robinson (76), there is a tendency to persist in the incorrect use of the name Banisteria. This tendency is [ 80 ] not always attributable to ignorance or uncertainty. It is frequently the result of a deliberate flouting of the International Rules of Botanical Nomenclature. Its continuation will not lead to standardization or to clarity but to the further beclouding of the exact identity of the malpighiaceous narcotics as well as to confusion in tax- onomic and floristic works. Ducke (19,20), in refusing to use the generic epithet Banisteriopsis, stated that his reason was an objection to the multiplication of names in botany. Macbride (48) not only chose to use Banisteria; he made the new combina- tion Banisteria inebrians, attributing it erroneously*® to Morton who has been most outspoken against this use of the name Banisteria. And recently Baldwin, in an article on the chromosomes of Banisteriopsis Caapi (5) chose to use the epithet Banisteria. Non-botanical writers, faced with this lack of standard- ization in technical papers, often continue to use Bani- steria instead of Banisteriopsis in connection with the narcotic species under discussion (21,380,382, 46,51,62,68, 64,91). An increasing number of technical writers, neverthe- less, are employing the correct generic epithet (12,28, 33, 34,65,85,93). V Although little new information has been published since 1981, a study of material preserved in our herbaria and museums, and recent field observations and collec- tions by several plant explorers have contributed other data which should be made available. It would seem to be especially necessary to do this, since, as the foregoing literature review has shown, little if anything of an ac- 3 ** Ranisteria inebrians Morton, Journ. Wash. Acad. Sci. 21: 485 (1931).”” curate nature may be looked for from the usual type of anthropological field investigation. In utilizing data found on herbarium labels, caution must be taken. Fora field botanist might jot down ‘‘used as a narcotic’’ on the basis of information supplied by a native and without sufficient checking or without himself having seen the plant thus employed. Nevertheless, such data may be of great value as an orientation in problems as complex as the one at hand. A further drawback—and a serious one—to the use of sterile specimens for botanical identification is the diffi- culty, in a genus of such vegetatively similar species, of arriving at a satisfactory determination. Usually, all we have to work with in studying the mal- pighiaceous narcotics is sterile material. The cultivated plants seem rarely or never to flower (probably because of constant cutting back), and the forest lianas blossom sporadically and are seldom found in flower by collectors. In one hundred years, for example, we have only one flowering collection of Banisteriopsis Caapi from the field (that is, excluding plants brought to flower in experi- ment stations or botanical gardens), and that is the type collection made by Spruce. My determinations of most of the older specimens concur closely with the identifica- tions made by Dr. C. V. Morton, who specialized in the Malpighiaceae. 1 have identified the numerous sterile herbarium specimens considered below with reserve, even though I have put in more than ten years of field study on the problem. As the result of prolonged study of liv- ing plants, one acquires some familiarity with certain of the variations which these species may show under dif- ferent natural habitats. But I must further point out that this long period of field study in itself has made me rather cautious about drawing categorical and far-reaching con- clusions from what herbarium material we have at hand [82 | at the present time. No one can feel more certain than I that we are just beginning our serious taxonomic studies of the complex of malpighiaceous narcotics. Perhaps the most revealing observation based on bo- tanical material to have been made since Spruce’s time has not hitherto been published. I was fortunate in dis- covering in the Museum of Economic Botany at the Royal Botanic Gardens at Kew several samples of the ‘“‘yajé plant,’’ comprising twigs and leaves (Econ. Mus. Kew 60-1913) from the Caqueta, Colombia. These were sent in 1918 by Perey EK. Wyndham, Esq., His Majes- ty’s Minister in Bogota. The leaves were determined by Dr. T. A. Sprague who wrote, in a communication at- tached to the specimens, that they represented species of Malpighiaceae (possibly Tetrapterys) and that ‘*the twigs no doubt belonged to one of them.’’ I have care- fully compared the leaves with specimens of the Mal- pighiaceae at Kew and would agree (insofar as it is pos- sible to determine such limited sterile material) with Sprague that they may be referable to Tetrapterys. This is the first time that a malpighiaceous genus other than Banisteriopsis has been mentioned as the source of a narcotic. With the Wyndham material at Kew, there are sev- eral communications concerning the specimens. It is ap- parent that Wyndham was moved to gather specimens of the yajé plant as the result of interest aroused by a newspaper article (3). This article spoke rather optimisti- cally of yajé as a cure for beri-beri, and it published a long letter from Dr. Rafael Zerda Bayén, a Colombian pharmacist, who put forth extraordinary claims concern- ing the telepathic properties of the vine. No botanical determination was offered. Sprague, in one of the com- munications with the Kew specimens, wrote: ‘*. . . Dr. R. Z. Bay6én says he has deposited specimens of the [ 38 ] ‘Bejuco de Yagé’ with the Ministry of Public Works and that he doesn’t think it desirable to send the speci- mens abroad! Apparently, the climber has not been identified. ”’ The collection Killip & Smith 27385, from Iquitos, Peru, represents Banisteriopsis Caapi and reports the following information: ‘‘Ayahuasca. Woody vine. Strong narcotic tea brewed from the leaves and stem produces fanciful dreams; also used as a cure for many diseases and as an intoxicating beverage.’’ Killip & Smith 29486 and 29825, from the same region, likewise repre- sent Banisteriopsis Caapi, but in connection with these two collections, the botanists made no notes as to their uses. Ducke 25258 is a flowering collection of Banisteri- opsis Caapi from a plant gathered in Amazonian Peru and cultivated in the Jardim Botdnico in Rio de Janeiro. Wilhams 3741 and 8224, also from the Iquitos area, like- wise are referable to Banisteriopsis Caapi, and are the basis of Williams’ report (108) that ‘‘the lower part of the stem and its leaves are crushed and boiled in water. When sufficiently triturated, the infusion is passed through a sieve to remove fibrous material, and to the residue water is added to render it drinkable. The resulting greenish- brown infusion has a disagreeable, bitter taste and is said to contain narcotic properties. ”’ From this wealth of material, we may judge that Ban- isteriopsis Caapi is rather commonly cultivated in Ama- zonian Peru. Still another collection, Seibert 2173 (also from the Amazonian basin of Peru but from the Rio Tahuamant in the Madre de Dios, far from the Iquitos area) has been determined as Banisteriopsis Caapi, even though its leaves are slightly atypical, due possibly to the fact that the plant was a wild liana in the jungle and not a cultivated shrub in open gardens. The collector reported: ‘‘Ayahuasca. Liana in forest. Infusion taken [ 84 ] internally and preparation from bark produces illusions. Used by Indians. Narcotic. ”’ It is from the Amazonian regions of Colombia, how- ever, that the greatest number of collections of the mal- pighiaceous drugs has recently been taken. Klug 1971 and Cuatrecasas 10597, both from the Comisaria del Putumayo, represent apparently Banisteriopsis Rusby- ana, 2 species usually characterized by large leaves. The former collection, from the village of Umbria, is reported to be called ‘‘chagro-panga or oco-yagé’’ ; the latter, from near Puerto Ospina on the Rio Putumayo, is known amongst the natives as yageuco (undoubtedly the same as oco-yagé) and is reputedly cultivated by the Kofan Indians who use the leaves in the preparation of the nar- cotic yajé. In 19538, I collected what appears to be Ban- isteriopsis Rusbyana near Mocoa, capital town of the Putumayo, under the name chagropanga; the leaves of this plant are reputedly used together with the pounded bark of B. inebrians in the preparation of the drink yaye. In 1931, Klug discovered in Umbria what turned out to be an undescribed species of Banisteriopsis. Described by Morton (58) as Banisteriopsis inebrians, this forest liana, which grows toa length of ninety feet, goes under the epithet of yagé del monte amongst the Ingano In- dians of the Putumayo. Klug collected ample botanical material of the yagé del monte and observed (58): ‘‘One of the most interesting plants found in the region of the upper courses of the Rios Putumayo and Caqueta is the yagé. The Indians make a beverage of either the wild or cultivated yagé, boiling it in a large earthenware vessel an entire day, until there is formed a sort of liquid, like the syrup of sugar cane. They add to the yagé the leaves and the young shoots of the branches of the oco-yagé or chagro-panga (No. 1971), and it is the addition of this plant which produces the ‘bluish aureole’ of their visions. *’ [ 35 | More recently, Banisteriopsis inebrians has been re- ported as the source of a narcotic in the same region. Cuatrecasas 10598, collected near Puerto Ospina, appar- ently represents this species and is cultivated under the name yagé by the Indians as the ‘‘principal ingredient”’ of the narcotic drink, the unboiled stem (‘‘tallo crudo’’) being employed. Likewise, Cuatrecasas 11061 was taken from a plant cultivated by the Kofiin Indians of the nearby Rio Sucumbios. In 1942, I found the same In- dians using cultivated Banisteriopsis inebrians, called oo- Jd in the Kofan language, as the source of a narcotic drink at Puerto Ospina and on the Sucumbios, and I ex- perimented in both localities with the intoxicant prepared from vines from which the collections Schultes 3452 (from a cultivated plant) and 3474 (from a wild liana) were made. The collection Schultes 3346, likewise apparently referable to Banisteriopsis inebrians, was taken from a vine pointed out by the Ingano Indians of Puerto Limoén on the nearby Rio Caquetaé as the plant from which, without admixture, they prepare their yajé. From Mocoa, capital town of the Putumayo and centre of the Inganos, still other collections of Banisteriopsis inebrians were made (Schultes & Smith 3037; Schultes & Cabrera 19113) with the field annotations that they were called ayahuasca or yajé and bejuco de oro (‘‘golden vine’’), that they were narcotic and that the leaves were used as a strong pur- gative. I have found Banisteriopsis inebrians to be used with and without the admixture of any other species of the genus, but decoctions of this species had marked nar- cotic effects each time I drank them, whether or not any admixtures had been used. What is probably Banisteriopsis quitensis was reported by Klug (Klug 1934) as yagé cultivado (58) in the Um- bria region. Later, Cuatrecasas (Cuatrecasas 10599) noted that the Kofiins near Puerto Ospina grow this spe- [ 36 | cies and use it in the preparation of the intoxicating yajé drink. He stated that the plant material is boiled to pre- pare the beverage. Notes with the collection Garefa- Barriga 4634a, which has been identified as probably referable to this same species, indicate that the Indians who live between Mocoa and Umbria (Inganos) call the vine yajé and prepare an intoxicant from it, with the ad- mixture of two other plants. Garcia-Barriga (25) stated that Banisteriopsis quitensis is ‘‘cultivated near their dwellings in order to have it at hand during the rainy season’? and that ‘‘they make with the stems a drink or beverage which they call yaje.’’ One of the admixtures, according to Garcia-Barriga, is the amaranthaceous A/- ternanthera Lehmanii Hieron., locally known as borra- chera or chicha (both of which terms refer to ‘‘intoxi- cant’? in Spanish); the other admixture could not be determined. Alternanthera Lehmanii is said by Garcia- Barriga to be added also to native beers or chichas as a condiment to increase their intoxicating properties. A sterile collection which seems to represent Banisteriopsis quitensis (Pérez- A rbeldez 639) was stated to be used in the region of Florencia on the Rio Orteguaza in preparing an inebriating drink. Williams (103) has reported Ban- isteriopsis quitensis to be used in the same way as B. Caapt in Amazonian Peru, where it is gathered from both wild and cultivated plants. The Indians of the Comisaria del Vaupés in Amazon- ian Colombia have conserved many of their aboriginal customs, such as the use of the malpighiaceous narcotics. Unlike the natives of the Putumayo, at the eastern foot- hills of the Andes, who use species of Banisteriopsis in a concentrated decoction made by boiling the plant ma- terial, the Indians of the Vaupés prepare a cold-water infusion of the bark. Banisteriopsis quitensis and B. in- ebrians seems to be lacking in the flora of the Vaupés, [ 37 | but B. Caapi and what appears to represent B. Rusbyana are cultivated for use as intoxicants. The collections cited below are all sterile and identification is, therefore, not certain, but, in each case, they were taken from plant material which I know, from personal experimentation, to possess narcotic properties. The 'Taiwano and Kabuyari Indians of the Rio Kana- nari, an affluent of the middle course of the Apaporis, prepare their kaheé from two kinds of Banisteriopsis, both cultivated. One vine had no leaves at the time of the Baile de la Sabaleta, when yajé is drunk, but the stems obviously belonged to a species of Banisteriopsis. The other is represented by the collection Schultes & Cabrera 13156a and corresponds in all vegetative charac- ters with Banisteriopsis Caapi. Those Makunas who live along the Rio Popeyaca, also an affluent of the Apaporis, prepare the drink in two ways: either with one species of Banisteriopsis or with two species of this genus, both cultivated. The drink is called haheé by the Makunas. When it is made from one species, Banisteriopsis Caapi (represented by Schultes & Cabrera 15587) is employed: this species is known in Makuna by the name reé-ma. When two species are used in the preparation of the drink, Banisteriopsis Caapi is mixed with a vine called me-ne-ka-heé-ma (‘‘vine of ha-heé™’) and referable probably to B. Rusbyana (Schultes & Cabrera 15588). Of Banisteriopsis Caapi, only the rasped bark is employed, but of B. Rusbyana either the leaves or the bark or both are utilized. With these natives, I took yajé twice; once made with bark from Schultes & Cabrera 15587 and leaves from Schultes & Cabrera 15588, and once with bark of Schultes & Cabrera 15587 alone. Intoxication was induced in both eases, and I was unable to note that one preparation had dif- ferent or stronger effects than the other. Both of these [ 38 | preparations are taken by all men of the tribe during dances. When the medicine-man employs the narcotic for purposes of diagnosis or to enable him to work evil on others, he takes a strong preparation of Banisteriopsis Caapi alone. The Makunas of the Popeyaca report that, in difficult cases of diagnosis, the medicine man will add a few crushed leaves of a tree which is abundant along the flood-banks of the river: gway-ee-ga-mo-yoo-he-reé (‘‘tree of the gill of fishes’’). This tree, represented by Schultes & Cabrera 15556, has been determined by Dr. R. W. Woodson as probably Malouetia Tamaquarina. The leaves contain an abundance of sticky, white latex and, as the species belongs to the A pocynaceae, possibly are poisonous in large doses. Indeed, there are persistent reports in the upper Amazon that the bones of the pa- juil are, at the time when Malouectia Tamaquarina sets fruit (which this bird is said to eat), highly poisonous to dogs which may eat them (86). I have never witnessed the use of Malouetia and was not able to corroborate the report with those who understand the properties of me- dicinal plants either amongst the Makunas or neighbor- ing peoples. The Indians of the headwaters of the Rio Piraparana cultivate caapi in almost every plot around their houses. Schultes & Cabrera 17209 from a Barasana Indian site on the Cano Teemeefa represents Banisteriopsis Caapi. A cold-water infusion of the rasped bark of this plant to which was added water in which dried tobacco leaves (Nicotiana Tabacum 1.) had been steeped had highly narcotic effects. Banisteriopsis Caapi is apparently the commonest source of the narcotic caapi drink in adjacent parts of Brazil. It was from Ipanoré on the Rio Uaupés near the Colombian boundary that Spruce collected the type specimen of this species. A recent collection, Ducke 153, [ 89 | taken from a plant cultivated in Mandos from material gathered along the Rio Curicuriari, an affluent of the upper Rio Negro, represents Banisteriopsis Caapi. During a year’s stay in the upper Rio Negro and its affuents in Brazil in 1947, I heard, on several occasions, reference to two kinds of caapi. As has been pointed out in detail above, it was from this region that Spruce re- ported a second kind of caapi, known locally as caapi- pinima, which he suggested might be referable to the apocynaceous vine Prestonia amazonica (Hlaemadictyon amazonicum). And it was likewise in this region that Ikoch-Griinberg found that the Tukanos distinguish two species of caapi, for only one of which (Banisteriopsis Caapi) he ventured a determination. It was my good fortune in 1948 to be able to witness the preparation of and to take a narcotic drink amongst the nomadic Maku Indians of the Ira-Igarapé, an affluent of the Rio Tikié which flows into the Rio Negro below Ipanoré. Specimens (Schultes & Lopez 10184) taken from the flowering vine, from the bark of which a cold-water infusion was made without the admixture of other plants, were found to represent an undescribed species of a mal- pighiaceous genus allied to Banisteriopsis— Tetrapterys methystica R. K. Schultes (87), The beverage prepared from Tetrapterys methystica has a yellowish hue, quite unlike the coffee-brown color characteristic of all prep- arations of Banisteriopsis which I have seen or taken. A small amount of stem material for chemical study which I gathered from this wild vine was lost in the overturning of my canoe, so nothing, unfortunately, can be stated concerning the chemical nature of the plant. ‘wo impor- tant points, however, should be emphasized in connection with this discovery: (1) T'etrapterys methystica may pos- sibly represent the second kind of caapi reported by Spruce and Koch-Griinberg, and it might be that the [ 40 | epithet caapi-pinima (‘‘painted caapi’’) alludes not to ‘‘painted leaves but to the unusual yellowish hue of the drink prepared from it; and (2) the tentative identifica- tion as ‘‘possibly Tetrapterys’’ of the sterile material sent to Kew by Wyndham from the Caqueta of Colom- bia would seem to be strengthened by the employment, even at such a distance, of a species of T'etrapterys in the elaboration of a drink with proven narcotic properties. SUMMARY 1. The narcotic drink known in the western Amazon- ian regions as caapi (Brazil and Colombia), yayé (Colom- bia) and ayahwasca (Ecuador, Peru and Bolivia) is made basically from the same or closely related plants of the Malpighiaceae. It is probable that the Indian names natema, nepe and pinde are synonymous with the three more commonly used names mentioned above. 2. The most widely employed species are members of the genus Banisteriopsis. The species most frequently used in Brazil, easternmost Colombia and much of the Amazon basin of Peru and Bolivia is Banisteriopsis Caapi, but B. Rusbyana seems also to be utilized in easternmost Colombia. In the westernmost fringe of the Amazon basin, along the Andean foothills of Colombia, Keuador and Peru, Banisteriopsis quitensis, B. mebrians and B. Rusbyana seem to be the species most preferred. 3. The closely related genus Tetrapterys is employed in Brazil and possibly in Colombia. The only species of Tetrapterys definitely identified as the source of the nar- cotic caapi is T. methystica. 4. Mascagnia psilophylla var. antifebrilis has been in- dicated as one possible source of ayahuasca, but this re- port is open to very serious doubt. [ 41 ] 5. The identification of yajé as a species of Aristolo- chia is definitely without foundation. 6. Prestonia amazonica (aemadictyon amazonicum) of the Apocynaceae has frequently been named as the source of yaqjyé and caapi. There is little or no reliable evidence that this vine is ever employed, at least as the prime ingredient, in preparing the narcotic drink. 7. The species of Banisteriopsis and Tetrapterys are known to be employed alone and to have narcotic prop- erties when thus used. Two species of Banisteriopsis may also occasionally be utilized together. 8. Non-malpighiaceous plants are known occasionally, but apparently not frequently, to be added as admixtures together with Banisteriopsis in some areas. Prestonia amazonica has been reported to be thus used in Brazil. The solanaceous Datura and the amaranthaceous A/fer- nanthera Lehmanii have been indicated as an added ingredient in Colombia. The apocynaceous Malouetia Tamaquarina may enter into the preparation of the nar- cotic in the Vaupés of Colombia, but this report could not be verified. Several unidentified plants have been mentioned as admixtures in Bolivia, Colombia and Peru. [ 42 ] EXPLANATION OF THE ILLUSTRATION Pirare VI. (Upper). Makuna Indian witeh-doctor gathering stems of Banisteriopsis Caapi (Spruce ex Griseb.) Morton (Schultes & Cabrera 15587) for preparation of narcotic drink. Rio Popeyaca, Com- isaria del Amazonas, Colombia. (Lower). Banisteriopsis inebrians Morton (Schultes & Cabrera 19113) cultivated in an Indian garden near Mocoa, Comisaria del Putumayo, Colombia. The rasped bark of this plant, which is referred to as yaje, is added to the leaves of Banisteriopsis Rusby- ana in the preparation of the narcotie drink. Photographs by Ricuary Evans Scuunres [ 44 ] | ri ATI EXPLANATION OF THE ILLUSTRATION Prare VIL. Banisrertopsts Ruspyana (Ndz. ) Morton (Schultes & Cabrera 19112) growing in secondary forest at Mocoa, Comisaria del Putumayo, Colom- bia. The leaves of this plant, which is called chag- ropanga, are added to the rasped bark of Banister- iopsis inebrians to prepare the yajé drink in the Mocoa area. Photograph by Ricuarp Evans Senurres A PLATE EXPLANATION OF THE ILLUSTRATION Prare VILL. Makuna witch-doctor under the in- fluence of caapi prepared in a cold-water infusion of the bark of Banisteriopsis Caapi (Spruce ex Griseb.) Morton (Schultes § Cabrera 15587) with no admixtures. Rio Popeyaca, Comisaria del Ama- zonas, Colombia. Photograph by Guitiermo Cano O, (48 | PLATE VIII 10. i. 12. LITERATURE CITED Albarracin, Leopoldo. 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[ 56 ] Cam BOTANICAL MUSEUM LEAFLETS HARVARD UNIVERSITY BRIDGE, MAssaAcHUsETTS, JUNE 26, 1957 THE EFFECTS OF CERTAIN GENES ON THE OUTER PISTILLATE GLUME OF MAIZE BY Watton C. GALINAT A comparison of the effects which various genes have on the structure of a given plant-organ should sometimes reveal which of the loci concerned were involved in the previous evolution of that organ and which of them might, during domestication of the species, contribute to its further evolution. he extreme susceptibility of the outer pistillate glume’ in the maize tribe (Maydeae) and the related tribe 4 ndropogoncae to evolutionary modifi- cations suggests that the genetic variation affecting de- velopment of this organ in maize (Zea Mays 1) may reflect some of its evolutionary changes. The changes in glume structure were drastic during evolution leading to the formation of the cupulate fruit case (Galinat, 1956). This organ evolved from a long foliaceous bract (d’rian- thus spp.) to a shorter, coriaceous structure marked by various types of sculpturing (Manisuris spp. ; Hackeloch- loa spp.) and, finally, to a highly lignified glume which is specialized in shape, texture and plane of divergence so as to bring about a closure of the narrow opening of a cupulate rachis-segment (Muchlaena and Tripsacum). ‘Further mention of the glumes of maize will refer only to the outer pistillate ones. Thus, we might expect that a study of the various glume phenotypes of modern maize would be revealing as to the past and possible further evolution of this glume dur- ing domestication. Three dominant or semi-dominant genes which affect principally the outer pistillate glume were included in this study. They are as follows: (1) Papyrescent (Pn gene on chromosome-7); (2) Tunicate (Zw gene on chromosome-4); (8) Vestigial glume (/¢ gene on chrom- osome-1). These genes were studied in heterozygous con- dition after they had been incorporated, by repeated backcrossing, into an isogenic background of a sweet corn (maize) inbred (Purdue 389). Their effects on the histological structure of this glume were examined in material prepared in the following manner: Normal (typical) and variant specimens were fixed at 18 days after pollination. ‘This material was later embedded in paraffin, according to the usual procedure, and then cut in cross-section at 12 uw. The sections were then stained by the standard safranin-fast-green technique. Finally, a projection apparatus was used in making tracings from comparable slides. Normal glumes. The glumes and other floral bracts of modern maize are so reduced that the mature grain emerges naked above them. Protection for the grain, which is provided by long floral bracts in most other grasses, is supplied by husks (modified leaf-sheaths) borne below the pistillate inflorescence (ear) on a condensed branch (shank). The normal glume has papery lateral wings (Plate LX, fig. 3) and the entire structure is homologous to a leaf- sheath with sheath-auricles. The glume-sheath, which is the counterpart of the leaf-sheath, may be coriaceous in texture (in many South American varieties) or, more commonly, it may resemble its counterpart in teosinte 58 | in being corneous or indurate (in teosinte-contaminated varieties from North and Central America). The degree of this induration has been used as an estimate of teosinte- introgression in archaeological maize, after the reliability of the method has been established (Galinat et a/, 1956). Internally, the mesophyll of the glume is divided into definite specialized regions. The induration is confined to a region of smaller cells extending from the outer epi- dermis to a line delimited by a row of vascular bundles about midway through a cross section (Plate X, fig. 8). As indicated by strong safranin staining, these smaller indurated cells are characterized by an accumulation of lignin in the secondary walls. The remainder of the glume, extending to the inner epidermis, consists of large parenchyma-cells. During final maturation of the glume, this parenchymatous tissue collapses from desiccation. The resulting shrinkage of the parenchyma toward the bundles causes an outline of the vascular system to be revealed as parallel ridges along the inner epidermis. The vascular arrangement of the normal glume differs slightly from that which occurs in a vegetative leaf- sheath. The glume has smaller bundles and, since this organ is determinative, they converge at the apex. In the leaf-sheath these principal bundles remain parallel as they continue on into the blade. As in the leaf-sheath, the parallel bundles are of two sizes with the larger ones alternating with one or two smaller ones. ‘The small cross connections which anastomose between the parallel bun- dles are confined to the apical region of the glume as compared to a distribution along the entire length of both the blade and sheath of the leat. The glume wings are non-vascular as are the ligules and sheath-auricles of the leaf-sheath. It seems that these vascular differences between glumes and leaf-sheaths are more in the nature of minor modifications. [ 59 | EXPLANATION OF THE ILLUSTRATION Prate IX. The outer pistillate glume of maize, as effected by various genes. Left and right speci- mens show dorsal and ventral views. 1, papyres- cent (Pn); 2, tunicate (Tu); 3, normal; 4, ves- tigial glume (Vg). Twice natural size. Photograph by Frank Wuire [ 60 | IX PLATE EXPLANATION OF THE ILLUSTRATION Prare X. Projection tracings of comparable cross- sectional views of the outer pistillate glume as this organ is affected by certain genes. In each figure the outer (abaxial ) epidermis is uppermost. 1,papy- rescent (Pn); 2, tunicate (7'w): 3, normal; 4, vestigial glume (J/g). About 80 times natural size. Drawn by Wavron C. GALinat Ba e westecte ge 2240 stlyanseeg > ABET | Weta’ o As in the leaf, stomata occur along both epidermal surfaces. In the glume, however, they are vestigial and obscure along the outer epidermis. Such reduction may result from the extreme lignification of the underlying tissue. Rather well-developed stomata are found on the inner surface directly opposite the larger vascular bun- dles. The fact that the maize-glume has stomata and yet does not have an opportunity to function in photosyn- thesis, seems to emphasize the leaf-like nature of this organ. The outer epidermis has a glabrous cuticle and may acquire a brown, red or purple coloration from the action of certain genes. Epidermal pubescence is greatly re- duced, except in the lateral wings. Papyrescent glumes. This dominant mutant is charac- terized by glumes which upon final maturation become papery and similar in texture to that which occurs in lem- mas, paleas and lateral wings of normal glumes, as well as in the ligules of normal leaves (Plate LX, fig. 1). These papyrescent glumes are usually slightly longer than the mature grain. A condition which is superficially similar to the papyrescent character develops in normal ears which are poorly matured. This simulation results from the protrusion of the lemmas, paleas and papery wings from normal glumes above chaffy or small grains. The papyrescent character is likewise similar or iden- tical with the ‘‘palee sviluppate’’ of Bonvicini (1932) and apparently with the recessive ‘‘semivestidos’’ of Andrés (1950). Our character for papyrescent glumes may have been incorrectly identified by Andrés as a recessive fac- tor since the effects of this semi-dominant gene are not always readily apparent in the heterozygote. It has also been confused by Weatherwax (1954) with half-tunicate, from which, as will be pointed out later, it is histologi- cally distinct. Furthermore, we find a counterpart of the [ G4 | papyrescent condition in Sorghum, where it once led to the describing of a new species (Sorghum papyrascens Stapf), an error later corrected by Rangaswami and Panduranga (1986) who correctly identified it as a single gene mutant characterized by defective glume develop- ment. Papery glumes have also appeared in two X-ray induced mutants (seminudoides and subnudoides) of bar- ley (Scholz, 1956). In these two recessive mutants, the floral bracts fragmentize away from the grain during threshing. Ordinarily the floral bracts of barley remain about the grain following threshing, except in the mutant named nudoides. The differences in the terminology of Sholz and Andrés, as applied to their particular mutants, refer to differences in the normal condition of the glumes in barley and maize. The term ‘‘seminudoides”’ (half- nude) applies to the barley mutant because normally the threshed barley grain is completely covered, while ‘‘semi- vestidos”’ (half-clothed) is better for the maize mutant because normally the maize grain is exposed. Papyrescent glumes in maize consist largely of non- specialized parenchyma-cells which, at 18 days from pol- lination, are large and fleshy, being swollen with water (Plate X, fig. 1). During final maturation of the ear, dessication causes these parenchymatous glumes to shrink to athin, almost transparent condition with the vascular bundles becoming prominent ridges (Plate LX, fig. 1). There is little contraction in length and width, however, and the glumes remain partially covering the surface of the mature grain. Finally, they become papery and brit- tle and are distinctly different from those glumes which protect the caryopses of other grasses. The epidermal layers of the papyrescent glume ap- proach those of the normal glume in regard to cell size, but differ from normal in being less lignified and in lack- ing stomata. Pubescence is usually confined to the mar- [ 65 ] ginal part of the glume. The vascular bundles are es- sentially normal in structure, although they are not so closely spaced. The succulent nature of immature papyrescent glumes often encourages the destructive activities of ear-rotting fungi. This suggests that a hardening of these glumes, such as is caused by teosinte introgression, may promote resistance to pathogenic fungi. The gene which produces this defective character has been previously designated as pseudopod (Pp) (Galinat and Mangelsdorf, 1955), but it now seems more desirable to change its name and symbol to papyrescent (Pn) (Galinat and Mangelsdorf, 1957). This change will call attention to its final papery character and its similarity to the “‘papyrascens’’ character of Sorghum, and will avoid confusion with the symbols for heterozygous peri- ‘arp color (Pp) as well. Our Px gene was originally iso- lated from a Peruvian variety of maize (Mangelsdorf, 1948). It is located close to the bd (branched-silkless) gene near the end of the long arm of chromosome-7. Its linkage relations will be discussed in more detail in a future paper. Tunicate glumes. The highest tunicate allele (7'w) from the series of multiple-alleles at the T’u-tu locus was chosen in order to study, in accentuated form, the effects of genes at this locus. Weaker alleles at the tunicate locus have intermediate effects on the length, shape and texture of the glume (Mangelsdorf, 1948). Further dis- cussion of tunicate glumes will refer to the phenotype of the strongest tunicate allele (Plate LX, fig. 2). The glumes of tunicate maize are like those of most other grasses in being long enough to enclose the grain and in being foliaceous. They differ therefore, in length and texture from the pistillate glumes characteristic of the American Maydeae, including normal maize, and of | 66 Manisuris spp. in the Andropogoneae. In the evolution- ary sequence leading to formation of the cupulate fruit case, tunicate glumes of maize are similar to the glumes of Mlyonurus tripsacoides (Galinat, 1956). Anatomically, tunicate glumes have all of the charac- teristics of a typical leaf-sheath, but on a reduced scale (Plate X, fig. 2). The close similarities of these organs may be enumerated as follows: (1) The inner (adaxial) and outer (abaxial) surfaces of tunicate glumes and leaf- sheaths are about equally pubescent with soft trichomes. This condition differs from that of the leaf-blade where pubescence is located chiefly on the upper (adaxial) epi- dermis and from that of the normal glume where the trichomes are confined to the lateral wings. (2) Both tunicate glumes and leaves (including sheath and blade) are herbaceous in texture because their mesophyll lacks the region of lignified cells which characterizes the nor- mal glume. (3) Stomatal development in tunicate glumes resembles that of the leaf-sheath and differs from the condition found in the normal glume. ‘Tunicate glumes and leaf-sheaths have well-developed stomata on both surfaces although they are more frequent and functional on the outer epidermis. As noted previously, if well- developed stomata occur in the normal glume, they are located on the inner surface. (4) The vascular system of tunicate glumes is more similar to the venation of leaves than it is to that of normal glumes. In both tunicate glumes and leaves, the anastomosing venation connects the principle bundles at irregular intervals along their entire length rather than just in the distal region as in the normal glume. (5) The structure of tunicate bundles is frequently identical to that of leaf bundles in contrast to the more reduced veins of normal glumes (Plate X, figs. 2 and 3). All of these anatomical comparisons sup- port the contention that the tunicate locus controls evo- [ 67] lution from the primitive foliaceous state which is typi- cal of glumes in general to the highly specialized glumes of modern maize. Vestigial glumes. The vestigial glume (Vg) mutant re- ported by Sprague (1989) is of particular morphological and agronomic significance. Originally this naked- flowered character was thought to be of no economic value because it was associated with pollen-blasting tas- sels. The discovery of modifying genes which usually permit abundant pollen production in Vg tassels, aroused interest in the utilization of the Ve character to improve the structure of the sweet corn (maize) ear (Galinat, 1951, 1953). It became apparent, however, that in order to insure pollen-production in inbred lines grown under various environmental conditions, it would be necessary to have tassels with glumes of normal length on Vg plants bearing glumeless ears. This desired Vg pheno- type became a reality after a certain weak tunicate allele derived from the race ‘‘Chapalote’’ was combined with the Vg gene (Galinat, 1955, 1957). As a result, the agronomic development and testing of sweet corn hybrids with this new Vg phenotype is now in progress. Vg glumes deviate from normal in the opposite direc- tion from the deviation of tunicate glumes (Plates [X and X, figs. 2, 3, 4). In comparison to normal ones, Vg glumes are thicker, shorter and have increased lignifica- tion, while 7’ glumes are thinner, longer and _ have re- duced lignification (lacking the lignified region in the mesophyll). The progressive thickening of glumes in this series of three types (7'u; normal; Vg) appears to result largely from the development and proliferation of many small lignified cells in a region just under the ab- axial epidermis. The smaller the cell in this region, the thicker the deposit of lignin in the secondary wall (Plate X1). [ 68 | The accumulation of this lignin is known to be second- ary, in that it occurs after the glumes have already at- tained optimum length and during the development of the caryopsis. If fertilization does not occur, then the glumes remain non-lignified and finally become papery. If only a portion of the ear is fertilized, then only those glumes which are either directly associated with fertilized ovaries or indirectly associated by being adjacent to spike- lets with fertilized ovaries will become lignified. This im- mediate effect of fertilization in stimulating lignification of maternal tissue occurs also in teosinte and T’ripsacum. Also the cupule of maize, as well as its counterpart in these close relatives, is included in the same metaxenial phenomenon. Certain features of the vestigial glume syndrome seem to be correlated with morphological homologies. The effect of the Vg gene in reducing glume size appears to be accomplished largely by suppression of the lateral wings (probably sheath-auricles), with the remaining por- tion representing a small, highly lignified sheath (com- pare figs. 8 and 4in Plate IX). In addition to the lateral wings of the glumes, the /’2 gene also causes, under cer- tain conditions, a reduction of the lateral wings of the cupules and the ligules of the leaves; this last-named effect was noted first by Laughnan (1956). Glume wings and ligules have at least two features in common in that both of them are papyraceous in texture and are situated as erect prolongations of the sheath, or its homologue, adjacent to the actual or theoretical insertion-point of the blade. Furthermore, these structures appear to be homologous, as is suggested by the vegetative leaves of certain grasses (4 mmophila arundinacea) with blades that are narrower than their sheaths. In such grasses, the lateral portions of the ‘‘ligule’’ elongate and develop as sheath-auricles with the same textural characters as the | G9 | EXPLANATION OF THE ILLUSTRATION PLare XI, A few contrasting cell types from a cross-sectional view of the outer pistillate glume of an ear with the vestigial glume character. Note that the smaller cells have a higher degree of lig- nification in the secondary walls (the stippled area). About 500 times natural size. Drawn by Warron C. GaLinat [ 70 ] | EG PLATE sheath, whereas the portion which is adjacent to the blade is reduced and papery. A similar phenomenon occurs in the lemmas of many grasses (4vena and Bromus) where the blade is modified into a narrow awn. The effect of the /¥@ gene in reducing the size of the cupule wings is not necessarily comparable to its effects on either glume wings or ligules. Rather the small size of Ie spikelets may decrease their capacity to depress the adjacent rachis-internode during youth and thereby result in shallow cupules. Discussion The features of Tu and V2 glumes differ from normal in Opposite ways and in doing so they reflect glume-ty pes involved in the early evolution of maize and T'ripsacum in the American Jaydeae. Teosinte is thought to have developed later from the hybridization of maize and Tripsacum (Mangelsdort and Reeves, 1939). The folia- ceous character of tunicate glumes is typical of the An- dropogonaceous grasses (as in HNlyonurus tripsacoides). Also it is a starting point from which the glumes of maize, Tripsacum, and possibly Manisuris could have evolved (Galinat, 1956). In the differentiation of T?ip- sacum and Manisuris, the glumes underwent reduction and lignification or sculpturing while the paired grain- bearing spikelets were reduced to singleness. Maize evolved in a separate direction which was controlled largely by mutation at the Z'w-tw locus (Mangelsdorf, 1948). Nevertheless, a latent ability to evolve a Tripsa- coid-type of outer glume might have been retained in modern maize and finally expressed as the /g mutant. We conclude that in regard to this series (7; normal; Vg), the structure of tunicate glumes indicates that the Tu-tu locus may have been involved in previous evolu- tion of the maize glume and that the mutation to the vestigial glume condition demonstrates a potential in maize to evolve a Tripsacoid-type of glume. The evolutionary changes resulting in shorter glumes during the domestication of maize could hardly have in- volved mutation at the ’g—vg locus. The Vg gene is a rare type of dominant mutation which seems to be of recent origin. Furthermore, it is not known to be part of a multiple-allelic series such as has been reported for the Tu-tu locus. There is also strong morphological evi- dence that the Vg gene has not been active in maize evolution. If such V’g activity had occurred, then it would be revealed by a characteristic reduction of the lateral wings of glumes. On the contrary, these wings are well-developed and papery in both modern and archaeological maize. In teosinte and Tripsacum, how- ever, the glume wings are somewhat reduced and highly lignified. In the series Tu; normal; Vg, decreases in glume- length are accompanied by increases in the thickness of the outer lignified zone. Since lignification of this outer zone of the glume occurs at the same time as kernel- development, there may be competition for the available energy. In the tunicate series of alleles (Mangelsdorf and Mangelsdorf, 1957) some of the energy conserved from shorter glumes is apparently diverted to increased grain production in spite of an increase in glume-lignification. Experiments are now in progress to determine the effect of the Vg gene on yield of mature grain. Pn glumes exemplify a condition which does not fit into this pattern of variation and, notwithstanding the suggestion of Andrés (1950), could scarcely have exer- cised a protective role at any stage in the evolution of maize. The only possible evolutionary counterpart of papyrescent glumes in other grasses appears to be repre- sented by a certain degenerate change found in several [73 | varieties of Sorghum (Rangaswami and Panduranga, 1936). A papery condition which is apparently similar to that of papyrescent maize has been produced by two X-ray induced mutations in barley (Scholz, 1956). ACKNOWLEDGMEN'TS During the preparation of the manuscript, many help- ful suggestions were made by Professor P. C. Mangels- dorf of Harvard University. Professor Angelo Bianchi of Pavia University helped in the preparation of some of the slides. LITERATURE CITED Andrés, J. M., 1950. Granos semivestidos, restos de un character ancestral del maiz. Rev. Argentina Agron. 17: 252-256. Bonvicini, M., 1932. Sulla ereditarieta di una anomalia del maiz. L’Italia Agricola 69: 3-9. Galinat, W. C., 1951. Glumeless hybrid sweet corn. Jour. Hered. 42: 114-116. ——, 1953. Resistance to blasting of Vg tassels. Maize Gen. Coop. News Letter 27: 76. , 1955. Maize with vestigial-glume ears and normal tassels. Maize Gen. Coop. News Letter 29: 27. ——, 1956. Evolution leading to the formation of the cupulate fruit case in the American Maydeae. Bot. Mus. Leafl. Harvard Univ. 17: 217-239. ———, 1957. The effect of weak tunicate alleles on the expression of the Vg gene. Maize Gen. Coop. News Letter 31: 68. Galinat, W. C., and P. C. Mangelsdorf, 1955. Pseudopod, a possible allele of vestigial glume. Maize Gen. Coop. News Letter 29: 26. ——, 1957. Papyrescent maize. Maize Gen. Coop. News Letter 31: 67. Galinat, W. C., P. C. Mangelsdorf, and L. Pierson, 1956. Estimates of teosinte introgression in archaeological maize. Bot. Mus. Leafl. Harvard Univ. 17: 101-124. Laughnan, J. R., 1956. Effect of Vg on development of the ligule. Maize Gen. Coop. News Letter 30: 67. Mangelsdorf, P. C., 1948. The role of pod corn in the origin and evo- lution of maize. Ann. Mo. Bot. Gard. 35: 377-398. and R. G. Reeves, 1939. The origin of Indian corn and its rel- atives. Texas Agric. Exper. Sta. Bull. 574. Mangelsdorf, P. C., and H. P. Mangelsdorf, 1957. Genotypes in- volving the J'u-tu locus compared in isogenic stocks, Maize Gen. Coop. News Letter $1: 65-67. 75 | Rangaswami Ayyanger, G. N., and V. Panduranga Rao, 1986. Sor- ghum papyrascens Stapf. Jour. Indian Bot. Soc. 15: 139-142. Scholz, F., 1956. Mutationsversuche an Kulturpflanzen V. Die Vere- bung zweier sich variabel manifestierender Ubergangmerkmale von bespelzter zu nackter Gerste bie réntgeninduzierten Mutanten. Die Kulturpflanze 4: 228-246. Sprague, G. F,, 1939. Heritable characters in maize 50—Vestigial Glume. Jour. Hered. 30: 143-145, Weatherwax, P., 1954. Indian corn in old America. The Macmillan Co., New York, 253 pp. BOTANICAL MUSEUM LEAFLETS HARVARD UNIVERSITY CAMBRIDGE, MAssACHUSETTS, JANUARY 27, 1958 VoL. 18, No. 3 ORCHIDACEAE NEOTROPICALES IV NoreEs ON ‘THE GENUS Caularthron Rar. BY Ricuarp Evans SCHULTES CAULARTHRON was recognized by Rafinesque as a generic concept distinct from EMpidendrum as early as 1836. It was treated as aspecial section of Mpidendrum by Lind- ley in 1841 under the name Diacrium. Section Diacrium was raised to generic status by Bentham in 1881. This is the name which has been applied to the genus by most subsequent orchidologists. Were the genus large or had it become important in horticulture, the resumption of an earlier and unfamiliar name might be unfortunate; and the best procedure might be an attempt to include Diacrium amongst the officially conserved generic names. I believe that this procedure should be reserved for names of large genera of great economic or horticultural importance, where the abandonment of a long and well established epithet would lead to extreme confusion. Therefore, | reeommend the substitution of Caularthron for Diacrium. Our herbaria have not had abundant material of Cau/- arthron, nor does the genus appear to have become a wide favorite in horticulture. In great part owing to these circumstances, it has not been so thoroughly understood as we might wish. Recent collections of Caularthron in Middle America, northern South America and Trinidad [ 77 ] and ‘Tobago have added somewhat to our knowledge of the range and variability of the genus, but there still re- mains much to clarify. In preparing the orchid section for the Fora of Trini- dad and Tobago, 1 have had to consider critically the concepts which have been known as Diacrium bicornutum and Diacrium indivisum. A study of the available her- barium specimens and of the very superior material pre- served in alcohol and sent in recently by Dr. Wilbur G. Downs and Dr. T. H. G. Aitken of Port-of-Spain, Trin- idad, made it early apparent that the fundamental prob- lems involved could not satisfactorily be handled without an examination of the generic concept as a whole. The present paper embodies the results of that examination. I wish to thank the officials of the following herbaria for making available material entrusted to their care: Reichenbach Herbarium (in the Naturhistorisches Mu- seum in Vienna); Gray Herbarium; Royal Botanic Gar- dens at Kew; New York Botanical Garden: United States National Herbarium; Chicago Natural History Museum and Missouri Botanical Garden. Material from these herbaria has supplemented the large collection of Caularthron preserved in the Orchid Herbarium of Oakes Ames of the Botanical Museum of Harvard University. It is, furthermore, a pleasure for me to thank Mr. G. C. K. Dunsterville for kind permission to reproduce two carefully executed drawings prepared for his forthcoming book of illustrations of Venezuelan orchids. When Bentham described Mpidendrum bicornutum in 1834 on the basis of material from Trinidad, he stated that he had consulted Lindley concerning its generic status, and had received the opinion that it ‘‘is certainly anew species; but I think it cannot be separated from Kpidendrum. The only distinction between it and that Genus consists in the labellum being distinct from the [78 | column: but you will find various degrees of separation between those parts. . .”’ in several species **. . . which nobody can doubt are genuine Lpidendra... Should you, however, be of opinion that it nevertheless must form a new Genus, its character will have to depend upon the large size of the peta/s and the slight adhesion of the sepals at their base.*” Seven years later, Lindley erected his subgeneric concept Diacrium on the basis of Epidendrum bicornutum. 'Then, in 1881, when he raised Diacrium to generic status, Bentham wrote that “‘the peculiar bicornute labellum, neither adnate to nor paral- lel with the column, gives the flower a very different aspect from that of true species of Hpidendrum and can not be included in them without doing violence to the generic character.’ Having now at hand material from a wide geographic range—all with certain diagnostic characters which show no variation—I believe that the best interests of orchid classification may be served by keeping the concept dis- tinct on a generic level. It is obviously very closely allied to Mpidendrum, and some of the differences used to sep- arate it may appear to be superficial. Nevertheless, it would seem that they represent perhaps a definite tan- gential evolutionary trend which ought to be recognized. There can be no doubt that Caularthron should be used as the name for this generic concept in preference to Diacrium. There is a widespread aversion to the accept- ance of many names proposed by Rafinesque. But I am sure that most botanists will agree with Merrill (Merrill, E.D.: ‘‘Index Rafinesquianus’’ (1949) 26, 29) that “*... where a new Rafinesque name was based wholly on a previously described or illustrated species of some other author, all we have to do to understand the application of the Rafinesque name is to determine the status of the originally described form . . . Thus it seems to be a logi- [ 79 | cal course to follow to continue to select the few sound grains from the overwhelming amount of chaff in the Rafinesque technical botanical papers, even if, occasion- ally, some more or less universally used generic or speci- fic name might fall before those proposed by Rafinesque at earlier dates. ”’ No greater precision could be desired than that which we find in Rafinesque’s description of Caularthron as a new genus. He not only published a very adequate de- scription which makes definite references to key morpho- logical characters separating the concept from Lpiden- drum, but, in a day when few botanists even mentioned what we now call types, he named the concept on which he was basing Cauwlarthron by citing Hooker’s Epiden- drum bicornutum and referring to its place of publication. The later name for this same generic concept, Diacrium, was likewise based on H’pidendrum bicornutum. The fact that, in second place under his generic description, Rafinesque made the new name Caularthron umbellatum, citing as basis for it Mpidendrum stenopetalum Hook., does not militate against the wisdom or the necessity of accepting as valid his generic name, especially so since his generic description is obviously based on Kpidendrum bicornutum and not on the very distinet 22. stenopetalum. Caularthron Rafinesque F 1. Tellur. 2 (1836 [1837] ) 40, pro parte. E’pidendrum Linnaeus sect. Diacrium Lindley in Hooker Journ. Bot. 3 (1841) 81; Bot. Reg. 81 (1845) Mise. 23; Fol. Orch. (1853) Epidendrum 8; Reichen- bach fil. in Walpers Ann. Bot. 6 (1862) 345. Miacrium (Lind).) Bentham in Journ. Linn. Soe. 18 (1881) 812; Bentham & Hooker fil. Gen. Plant. 3 (1883) 526; Hemsley Biol. Centr.-Am. Bot. 3 (1883) 221; Warner & Williams, Orch. Alb. 4 (1885) t. 157; [ 80 | Rolfe in Gard. Chron. 2, ser. 8 (1887) 44; Pfitzer in Engler & Prantl Nat. Pflanzenfam. II, 6 (1888) 146; Veitch, Man. Orch. Plant. 6 (1890) 78: L’Orchidoph. (1891) 878; Rolfe in Lindenia 7 (1891) 19; Cogniaux in Martius Fl. Bras. 8, pt. 5 (1901) 186; Stein, Orch- ideenb. (1892) 214; Bois, Les Orch. (1898) 74: Ner- chove, Livre des Orch. (1894) 264; Linden, Orch. Exot. (1894) 751; Cogniaux in Urban Symb. Antill. 6 (1910) 588: Schlechter, Die Orchideen (1915) 214: Ames ex Standley in Field Mus. Nat. Hist. Publ. 391 (1937) 210: L. O. Williams in Ann. Mo. Bot. Gard. 33 (1946) 878; Hoehne, Icon. Orch. Bras. (1949) 208 ; P. H. Allen in Orch. Journ. 2 (1953) 185; Ames & Correll in Fieldiana: Bot. 26 (1953) 405; Foldats in An. Univ. Centr. Venez. 34 (1953) 276. Caularthron may be distinguished from Lpidendrum on the basis of the characters set forth in the following key. A. Labellum vulgo ad columnam variabiliter adnatum et cum ea parallelum, supra numquam protuberantibus elevatis subtus ex- cavatis ornatum, Epidendrum Aa. Labellum a columna liberum, a columnae basi angulatim patens, supra protuberantibus duabus elevatis subtus excavatis ornatum Caularthron E\piphytic or semi-epiphytic herbs with fleshy pseudo- bulbose, solid or frequently insect-hollowed stems. Leaves few, borne at apex of pseudobulbs, rigidly sub- ‘arnose-coriaceous. Inflorescence terminal, simple, race- mose. Flowers few to numerous, showy, short-pedicel- late. Sepals free, spreading, subequal. Petals rather similar to sepals. Lip free and spreading from base of column, 3-lobed; lateral lobes conspicuously tooth-like ; mid-lobe triangular or triangular-lanceolate; disk raised between lateral lobes into 2 hollow, often horn-like cal- luses opening from below to form 2 conspicuous inden- [ 81 ] EXPLANATION OF THE ILLUSTRATION Pirate XIT. Cautarruron psicornutum (Hook. ) Rafinesque. Grown in the greenhouse of F. W. Hunnewell, Wellesley, Massachusetts. Photograph by Ross W. Baker Courtesy of American Orchid Society, Inc. [ 82 | PLATE NII EXPLANATION OF THE ILLUSTRATION Prare XII. Caurarruron prcornurum (Hook. ) Rafinesque. Drawing of Dunsterville 399 trom Ven- ezuela. Drawn by G. C.K. Dunstrervitir [ 84 | XIII PLATE tations or pits on lower surface of lip. Column short, with conspicuous fleshy wings; clinandrium oblique, obtuse. Anther terminal, operculate, incumbent, more or less globose, 2-celled; cells divided longitudinally. Pollinia 4, waxy, each with a granular-viscid appendage. Capsule ellipsoidal. Caularthron: from the Greek, meaning ‘‘jointed stem,’’ undoubtedly in reference to the persistent leaf- bases which lend to the elongated pseudobulbs the ap- pearance of being jointed. A genus occurring from Guatemala through Middle America, northern South America and Trinidad and Tobago. Two rather variable species are known. Key to the species of Caularthron A. Labellum 24-28 mm. longum, profunde trilobatum, lobis usuali- ter sinu conspicuo separatis; lobo mediano propie lanceolato vel elongato-lanceolato; lobis lateralibus ovato-oblongis. Folia ob- longa vel oblongo ligulata, 7-25 (plerumque 12-20) em. longa <1,5-4.2 (plerumque 3-4) em. lata. 1. Caularthron bicornutum Aa. Labellum 11-14 mm. longum, saepissime inconspicue trilobula- tum vel auriculatum vel aliquando subintegrum, lobis usualiter non sinu separatis; lobo mediano propie triangulari-ovato; lobis lateralibus parvis, vulgo auriformibus. Folia ligulato-lanceolata, 7-19 (plerumque 14-15) em. longa x 0.8-2.8 (plerumque 1-1.8) em. lata. 2. Caularthron bilamellatum ORIGINAL DESCRIPTION ! 138. Caularthron R. (Stem jointed) Diff. Epidendron. Label. libero, ad basi alato glanduloso. Col. libera bialata dentata, anthera termi- nalis 4 pollen. Caul. articulato, vaginato, bifolio, paucifloro—Habit very irregular. Types 2 Sp. 1. Caul. bicornutum. Epid. do Hook. b. m. 3332. Bulbo cauliformis, fol. rad. ligul. retusis, scapo paucifl. label. trilobo, medio lane. ae basi bicorne, petalis ellipt. acutis albis. Trin- idad... Caularthron bicornutum (Hook.) Rafinesque F. Tellur. 2 (1836 [1837] ) 41. Hpidendrum bicornutum Hooker in Bot. Mag. (1834) t. 833882. [ 86 | Diacrium bicornutum (Hook.) Bentham in Journ. Linn. Soc. 18 (1881) 3812. Diacrium amazonicum Schlechter in Beih. Bot. Cen- tralbl. 42, Abt. 2 (1925) 108; Pabst in Arqu. Bot. Est. Sado Paulo 3 (1955) 125, t. 316. CoMMON NAME: Virgin Mary; Virgin Orchid (Trin- idad and Tobago). Pseudobulbs long-cylindric, terete, 10-80 cm. long, 2-6 em. in diameter. Leaves 3-4, thick, oblong, usually quite obtuse to rounded, 6-20 (mostly 15-17) cm. long, 20-50 mm. wide. Flowers few to 20, 5.5-6 ecm. wide; pedicel (with ovary) 8-5 cm. long. Sepals broadly ovate- lanceolate, bluntly short-acuminate, 25-82 mm. long, 15-18 mm. wide. Petals ovate, usually clawed, acutish, 22-28 mm. long, 20-23 mm. wide, upper margin usually with a conspicuous notch. Lip fleshy, as long as petals but narrower, deeply 3-lobate, basally with triangular tooth on each side; lateral lobes elliptic-ovate, rounded; mid-lobe oblong, obtuse-acuminate; disk above with 2 erect, triangular, plate-like, obtuse, hollow projections near middle. Column 14-15 mm. long. The type of Caularthron bicornutum was collected in Trinidad by Bradford, who wrote on the label: ‘*This most beautiful species is found in the greatest abundance on the coast and on the adjacent islands at the Boca de Moros, Trinidad.—The rocks and trunks of decaying trees are in some places covered with it. This specimen was gathered on Gaspare Island March 12, 1846 on my return from an expedition to the coast of Venezuela. It flowers especially in the early part of the year from Jan- uary to April.” The culture of Caularthron bicornutum requires a rather warm greenhouse with high humidity (Warner, R. & B.S. Williams: Orch. Alb. 4 (1885) t. 157; Rolfe, | 87 | R. A.: Gard. Chron. 2 (1887) 44; Lindenia 7 (1891) 19; Linden, L.; Orch. Exot. (1894) 752). Much difficulty has sometimes been experienced in establishing the plant, as the pseudobulbs, although large and apparently rather tough, seem to damage easily. Once established, it seems to thrive well under cultivation, especially if not dis- turbed. ‘The most recent article dealing with its culture (Schairer, J. Fo: Bull. Am. Orch. Soc. 24 (1955) 106, t. p. 107) states: ‘‘Plants are grown potted firmly in brown osmunda. During autumn and winter, when the plants are in active growth, they require a warm spot (night minimum 63° F if possible), high humidity and good light (as much or more than most Cattleyas) with plenty of water at the roots, and they appreciate supple- mental feeding. At blooming time, they prefer a cooler spot and less light and moisture. They never require a severe rest period after blooming but are kept somewhat dry for a few months before new growth begins. They are native of the West Indies and the Guianas, where they often grow on bare rocks and tree trunks near the sea where they get plenty of moisture and cooling breezes during hot weather. Propagation is by division and they never propagate fast enough to supply the insistent de- mands of your friends. ”’ Caularthron bicornutum and C. bilamellatum occupy rather clearly defined geographic areas: the former is native to South America north of the Amazon River and to Trinidad and ‘Tobago, whereas the latter occurs in Middle America, along the northernmost rim of South America and in Trinidad. It is in Trinidad chiefly that the two species are contingent, but here there appears to be an ecological delimitation of the two species— Cau- larthron bicornutum forms a conspicuous element along and near the sea coast, while C. bilamellatum is known only from inland districts. [ 88 | There are anumber of minor morphological characters which serve to distinguish Caularthron bicornutum from C. bilamellatum. The former is, in general, a much larger and more robust plant than the latter and has flowers approximately twice as large. This difference is variable, and the specimens of Caularthron bicornutum from Brit- ish Guiana are notably smaller in habit and flowers than those from other parts of the range of the species. Caularthron bicornutum may be recognized at once through its deeply trilobate lip. The mid-lobe is charac- teristically lanceolate or elongate-lanceolate with the api- cal portion either long- or short-acuminate. The lateral lobes, which are ovate-oblong and either obtuse or sub- acute, are usually separated from the mid-lobe by a deep sinus. Caularthron bilamellatum, on the contrary, usually has an inconspicuously trilobulate or auriculate lip, the mid-lobe of which, characteristically triangular-ovate and acute, is not separated from the small, auriform lateral lobes by a recognizable sinus. ‘To be sure, several collec- tions are known from Trinidad which show a somewhat intermediate lobation of the lip, and here there may be evidence of hybridization. When the available material of these two species from their entire geographic ranges is taken into consideration, however, the significance of the shape of the lip as indicative of a possible evolutionary trend may be appreciated. A convenient character for separating Caularthron bicornutum trom C. bilamellatum—and a character which seems to have been overlooked—is found in the peculiar notching of the upper margin of the petals of C. becornu- tum. This margin, which faces the dorsal sepal, usually has one conspicuous notch situated one-third or one-half of the distance from the base of the petal. In some of the specimens, the notch is sharp and triangular; in others, it is less clearly defined. In all cases, however, [ 89 | the presence of this notch causes the marginal area of the petal to be somewhat ruffled or crumpled. I have very rarely observed anything similar to this condition in Caularthron bilamellatum. In the general shape of the hollow processes or ‘‘horn- like calluses*’ which arise from the disk of the lip and which have been used as the outstanding generic charac- ter, there is evident little variability from specimen to specimen, albeit some variation in relative size may be seen. Similarly, there is no significant morphological difference between these processes in Caularthron bicor- nutum and C. bilamellatum. Unlike the concept Caularthron bilamellatum, C. bicor- nutum does not have a large synonymy. This is due partly, perhaps, to the greater variability in the former than in the latter species. In 1925, Schlechter described Diacrium amazonicum from material collected in Brazil. The type material of Diacrium amazonicum is no longer extant, but a study of the type description convinces me that Schlechter specified no differences which, with the material at hand today, we could not easily accommodate in Caularthron bicornutum. Pabst (loc. cit.) determined a specimen (Froes 21541) from the Rio Negro in Amazonian Brazil as representing Diacrium amazonicum and published a diagnostic drawing of the floral parts. This drawing like- wise shows no character which could serve to distinguish the plant from Caularthron bicornutum. Type cottection: Cult. Hort. Wentwlorth], from Trinidad (Herb. Kew). Curtivarep: (Herb. Rehb. 898); Hort. Rucker, May 27, 1840 (Herb. Rehb. 899; Herb. Ames 69096a); ‘“‘from the type plant of Gard. Chron. 1887, pt. 2, p. 45, fig. 11,’ Hort. Kew, May 1887, R.A. Rolfe s.n. (Herb. Kew); Botanic Station, Grenada, W.I. ‘‘Orig- inally from Trinidad,’’ April 14, 1906, W. EH. Broadway s.n. (Herb. Ames 7906, 7907). | 90 | Brazit: Estado do Amazonas, Mandos, Spengler s.n. (Herb. Rehb. 896); Estado do Amazonas, Rio Negro, December 21, 1945, R. L. Frées 21541 (Record in Herb. Ames of specimen in Herb. Inst. Agron. Norte). Bririsn Guiana: *‘Flowers white, pink towards the apex, labellum pink with a yellow disk.’’ 1837, R. Schomburgk 429 (Herb. Kew; Herb. Field Mus. 1025283); Banks of Corentyne River near Crealla, September 1879, E. F. im Thurn s.n. (Herb. Kew); Essequibo River, December 1886, Jenman 3590 (Herb. Kew); Jenman 7761 (Herb. Kew); Hort. Kew, May 1889 (Herb. Kew); Mount Roraima, Autumn 1894, J.J. Quelch & F. McConnell 280 (Herb. Kew); C. F. Appun 657 (Herb. Kew); Cuyuni River, islet at the Akaio Falls. “‘Epiphyte at about 15 feet; roots in dense clusters. Fl. shallow, cup-shaped, like an Anthericum, pedicels mauve. Fl]. resupinate. Pets. and sep. pure white, with striations. Stele at base and all labellum spotted with purple. Humps of label. and depression between them and stele yel- low, spotted with purple. Label. otherwise white.’’ November 25, 1929, N. Y. Sandwith 685 (Herb. Kew). Cotompr1a: Comisaria del Vaupés, Rio Negro, El Castillo (San Fe- lipe). ‘‘Flowers white, sepals delicately pink-tinged. Lip yellow, spotted brown. Very fragrant.’? December 12, 1947, R. EF. Schultes & F. Lopes 9355a (Herb. Ames 67526, 67527). Tonaco: G. W. Meyer (?) s.n., December 1879 (Herb. Kew); Rox- borough Bay, Military Road, “growing on the stems of logwood tree,’”’ February 2, 1879, G. W. Meyer s.n. (Herb. Kew); Bacolet, ‘*on rocks and trees near the sea, flowers white with a few purplish spots, very fragrant,’ January 20, 1910, W. E. Broadway s.n. (Herb. Gray 4278); Rockley Vale, ““on trees, Virgin Mary,’’ February 20, 1913, W. E. Broadway s.n. (Herb. Gray 4277; U.S. Nat. Herb. 759445); Feb- ruary 19, 1932, D. Fairchild 2930 (U.S. Nat. Herb. 1625959; Herb. Ames 69057). Trinwwap: Borroughs s.n. (2) (Herb. Rehb. 896); Bradford 1846 Herb. [Hance 5332] Herb. Kew); Gasparee Island, ‘on rocks and trees overhanging the sea,’’ December 30, 1906, W. EF. Broadway s.n. (Herb. Ames 10085, 10086, 10087, 10088, 10089, 68642; Herb. Field Mus. 464601); Maraval, ‘‘on rocks and trees,’” February 8, 1911, W. E. Broadway s.n. (Herb. Kew; Herb. Ames 69058); Moruga sea shore, Lance Mettan. “‘On shrubs, trees and rocks.’* February 9, 1916, W. E. Broadway s.n. (Herb. Trin. 7595; Ames 69924); “‘On a tree,’ May 26, 1918, W. EF. Broadway s.n. (Herb. Ames 22062; Herb N. Y. Bot. Gard.); Manzanilla, ‘‘Flowers white. On a fallen tree,’? March 9, 1921, N. L. Britton & E. G. Britton 2173 (Herb. N.Y. Bot. Gard.; U.S. Nat. Herb. 1198102); Little Gasparee, April 4, 1921, N. L. Britton 2659 (Herb. N.Y. Bot. Gard.); Chacachacare, [ 91 | January 5, 1922, W. E. Broadway s.n. (Herb. Trin. 10566, 2 sheets); Herb. Ames 69925); St. Ann’s (Cult.) ‘‘wild on rocks and trees along sea shores,’’ March 8, 1923, W. E. Broadway s.n. (Herb. Field Mus. 549522); Between Balandra and Toco, “‘rocks by shore,’? February 2, 1926, W.G. Freeman s.n. (Herb. Trin. 11519); Balandra Bay, ‘‘on rocks and low trees near the sea shore. Virgin Orchid,’’? February 22, 1931, W. E. Broadway s.n. (U.S. Nat. Herb. 1519971: Herb. Mo. Bot. Gard. 1005390); “*On trees and rocks, sea shore districts: Vir- gin Orchid,’’ February 23, 1934, W. EF. Broadway s.n. (Herb. Ames 40385; Herb. Kew); Quinam Bay, St. Patrick, “‘On trunk of tree, fls. white with faintly speckled throat,’’? January 80, 1946, L. H. Bailey 121 (Herb. Ames 62541); Chacachacare Island, January 20, 1956 [Capt. Mendez] W. G. Downs & T. H. G. Aitken 15f (Herb. Ames Ale. Coll, 3187); Sangre Grande, Rio Grande Forest Tree Station, 14 mile from coast, March 7, 1956, W. G. Downs & T. H. G. Aitken 15g (Herb. Ames Ale. Coll. 3183); Chachachacare Island, scacoast, January 14, 1957 [A. S. Fenwick| W. G. Downs & T'. H. G. Aitken 15c (Herb. Ames Ale. Coll. $204a); Vega de Oropouche, 1 mile from sea, W.G. Downs & T. H. G. Aitken 15a (Herb. Ames Alc. Coll. 3206a). Venezurta: [Drawing of a flowering specimen] Carabobo, alt. 2500 ft., 1851 (Herb. Rehb. 890; Herb. Ames 69098); Paria Peninsula, Cariaquita, January 16-21, 1911, F. FE. Bond, T. 8. Gillin & S. Brown 40 (Herb. N.Y. Bot. Gard., U.S. Nat. Herb. 1189830); vicinity of Cristobal Colon, January 5—February 22, 1923, W. E. Broadway 337 (Herb. Gray 4275; Herb. N.Y. Bot. Gard. ; U.S. Nat. Herb. 1187925, 1197666); vicinity of Crist6ébal Colon, Avicana, January 5—February 22, 1923, W. FE. Broadway 616 (Herb. Gray 4276; U.S. Nat. Herb. 1197675); April 29, 1941, H. Pittier s.n. (Herb. Ames 68209); [Draw- ing of a flowering specimen] G. C. K. Dunsterville 399 | Arrigo R. s.n. |, Puerto Ayacucho (Herb. Garay 6358). Caularthron bilamellatum (?chb.f.) R.H. Schultes comb, nov. E’pidendrum bilamellatum Reichenbach fil. in Walpers Ann. Bot. 6 (1862) 845. E’pidendrum bigibberosum Reichenbach fil. loc. cit. 8 (1862) 346. Epidendrum indivisum Bradford ex Grisebach FI. Brit. W. Ind. Isl. (1864) 614. Diacrium bigibberosum (Rehb.f.) Hemsley in Godman & Salvin Biol. Centr.-Am., Bot. 8 (1883) 222. [ 92 | Diacrium bilamellatum (Rehb.f.) Hemsley in Godman Diacrium indwisum (Bradf. ex Griseb.) Broadway in Bull. Mise. Inform. Trinidad 2 (1895) 79. Diacrium bicornutum (Hook.) Bentham var. indivisum (Bradf. ex Griseb.) Cogniaux in Martius FI]. Bras. 38, pt. 5 (1898) 188. Diacrium venezuelanum Schlechter in Fedde Repert. Sp. Nov. Beih. 6 (1919) 41. Diacrium bilamellatum (Rchb.f) Hemsley var. Reich- enbachianum Schlechter loc. cit. 17 (1922) 47, in textu. Diacrium bwalvatulum Schlechter loc. cit. 19 (1923) 128. Pseudobulbs subcylindric to long-fusiform, terete, 5-23 cm. long, up to 4 cm. in diameter. Leaves 2-3, ligulate-lanceolate to linear-oblong, obtuse, 5-22 cm. (mostly more or less 15) em. long, 6-25 mm. wide. In- florescence erect, up to 15 em. long. Flowers white or white tinged with pink or lavender, few to numerous, up to about 8 em. wide; pedicel stout, (with ovary) 1.5-2.5 em. long. Bracts triangular, cucullate, acute, up to 5.5 mim. long. Sepals concave, elliptic-ovate, acute or some- times subobtuse and apiculate; dorsal sepal 12-17.5 mm. long, 6 mm. wide; lateral sepals 10-16 mm. long, 6-8 mm. wide. Petals short-clawed, elliptic-obovate to broadly obovate, acute or subobtuse, 10-16 mm. long, 6-9 mm. wide. Lip fleshy, as long as petals, subentire to more or less 38-lobulate; lateral lobes small and auricu- late, sometimes only tooth-like, usually without a sinus; mid-lobe triangular-ovate, apically short- or long-acute ; disk above with 2 large, erect, triangular, plate-like, ob- tuse, hollow projections. Column 8-11 mm. long. Cap- sule up to 2.8 em. long. The type of Hpidendrum bilamellatum was collected in ‘*Caracas’’ (referring probably to a very extensive area [ 93 ] EXPLANATION OF THE ILLUSTRATION Pirate XIV. CavuiarTuron pirametiatum (Rchbf.) R.E. Schultes. 1, flowering and fruiting plant, one third natural size. 2, flower, approximately natural size. 3, column and lip, side view, almost twice natural size. Drawn by Gorvon W. DiLLon [ 94 ] PuateE XIV in Venezuela and not specifically to the city), Venezuela, by Wagener and is preserved in the Reichenbach Her- barium on sheet No. 891. This sheet, which is labelled “Epid. bicornutum’’ has, pasted on a card attached to the sheet, three flowers from Fendler 2436, collected near Colonia ‘Tovar in Venezuela in 1856-1857. This Fendler material, representing Caularthron bilamellatum, has been given the number of Herb. Reichenbach 891a in order to distinguish it from the type or Wagener col- lection. There are also, pasted on the sheet with the type, two labels indicating that the collection was made by Linden in ‘‘Nouvelle Granade,’’ but since the original handwritten label is obviously the correct one, we must assume that, as so often happened, the printed Linden labels were glued to the sheet at a subsequent date and undoubtedly in error. An examination of the type material of Hpidendrum bigibberosum (Herb. Reichenbach 893, 894) and of Reich- enbach’s diagnostic sketches of the floral parts fails to uncover a single character which might serve to distin- guish this collection from the type of 2. bilamellatum. In his original description of the concept Hpidendrum bigibberosum, Reichenbach likewise failed to point out any differences. We, therefore, must reduce Mpidendrum bigibberosum to synonymy under Caularthron bilamella- tum. For some time, I have been undecided as to what the concept which has been known as Diacrium indivisum (Epidendrum indivisum) really represented. Bradford drew up a description based on one of his Trinidad col- lections, and this was published as Mpidendrum indivisum by Grisebach. It was transferred to Diacrium by Broad- way in 1895. Cogniaux maintained that the concept represented a variety of Diacrium bicornutum and made the necessary nomenclatural adjustment. In 1956, I pub- [ 96 ] lished a note indicating my belief that there seemed to be sufficient morphological evidence to maintain it [Dr- aerium indivisum| as specifically distinct from the only other Trinidad representative of the genus. Subsequent- ly, when Dr. Wilbur G. Downs sent me from Trinidad photographs and additional material of the two concepts of Caularthron known to grow on the island, I began to realize that a revision of the genus Diacrium was neces- sary before a clear understanding of the Trinidad material could be expected. The type specimen of EHpidendrum imdivisum is pre- served at Kew, together with Bradford’s handwritten description of the concept. The type bears the annota- tion ‘‘Herb. Hance 5334.’” On the same sheet with the type there are pasted two inflorescences from plants col- lected in Trinidad and flowered at Kew in May 1889. Rolfe has annotated this collection as ‘‘E/pidendrum bi- cornutum var. cleistogamic flowers=D. indivisuwm.”” According to Bradford’s manuscript description, the type has an undivided lip. In the published description, the lip of the type was stated to be ‘undivided or mi- nutely auricled above the base.”’ The apical part of the lip was described as ‘tacuminatum’”’ in the manuscript and ‘‘subulate-lanceolate’’ in Grisebach’s Fora. Untor- tunately, the type has, at the present time, only two buds and one imperfect flower. We know from later ma- terial, however, that, in Trinidad, this concept is often cleistogamous. It is possible that the type flowers may have been peloric. At least, we do know from the ma- terial now available from Trinidad that the lip is very rarely entire but is most often laterally auriculate at the base or inconspicuously bilobulate. In this, as in other respects, the concept does not depart from Caularthron bilamellatum, of which it is, consequently, here desig- nated as a synonym. [ 97 | EXPLANATION OF THE ILLUSTRATION Prare XV, CauLarruron BILAMELLATUM (Rehb f.) R. EF. Schultes. Drawing of Dunsterville 181 from Venezuela. Drawn by G. C. K. Dunsrervitte [ 98 | PLATE XV Schlechter’s Diacrium venezuelanum, a record of the type of which is preserved in the Ames Herbarium, shows no character which cannot fall easily within the variability of Caularthron bilamellatum. Diacrium bival- vatulum, likewise, presents, according to Schlechter’s original description, no differences of sufficient impor- tance for the maintenance of it as a distinct species. Dia- cerium venezuelanum and D. bivalvatulum are, therefore, placed in synonymy under Caularthron bilamellatum. Type cottection: Venezuela, ‘‘Caracas,”’ Wagener s.n. (Herb. Rehb. 891; Herb. Ames 69055). CoLompia: 1842, Sinclair s.n. (Herb. Kew). British Honpuras: W. A. Schipp S-482 (Herb. Ames 40443). Cosra Rica: [Drawing of a flowering specimen and diagnosis of flower] Punta Arenas, February 1909, 4. & C. Brade 1265 (Type of D. bivalvatulum) ; Provincia de Alajuela, El Coyolar, alt. about 240 m. “On tree; bulbs elongate, full of ants.’’ April 1-3, 1924, P. C. Standley 39982 (Herb. Ames 29865); Golfo Dulce, Playa Blanca, sea level, February 25, 1938, M. Valerio 553 (Herb. Field Mus. 893155). GuatemaLa: Cult. Hort. Shiller, from Guatemala, Skinner 1458 (Typr of FE. bigibberosum, Herb. Rehb. 893, 894; Herb. Ames 24081, 69091); Montufor Flats, February 27, 1939, M. W. Lewis 217 (Herb. Ames 69926); Departamento de Izabal, between Milla 49.5 and Cris- tina, alt. 65-70 m. ‘‘On limbs of tree along wooded margins of prairie. Pseudobulb terete, pale green. Leaves thick, coriaceous, rich olive- green. Stem olive-green with purple suffused above and at nodes. Pedicels purplish. Buds orchid-colored. 8 outer sepals delicate orchid without, white within. 2 petals white. Lip white on upper petal and stigma and white on beak with pale orchid color along edge or the column with more lavendar.’’ J. A. Steyermark 38389 (Herb. Field Mus. 1043049); Departamento de Izabal, Bay of Santo Tomas, be- tween Escobas and Santo Tomas, alt. sea level to2 m. April 13, 1940, J. A, Steyermark 39341 (Herb. Field Mus. 1041145; Herb. Ames 63752); Departamento de Izabal, shores of Lago Izab4l, opposite San Felipe; between San Felipe and mouth of Rio Juan Vicente, alt. 50 m. April 19, 1940, J. 4. Steyermark 39692 (Herb. Field Mus. 1035306). Panama: Santa Rita Trail, February 27, 1905, J. F. Cowell 160 (Herb. N.Y. Bot. Gard.); Cultivated from Canal Zone, Culebra, alt. 50-100 m., April 8, 1911, H. Pittier 3406 (Herb, Ames 21797); Pa- tifio, southern Darien, ‘‘on cliffs along the beach,’’ February 13, 1912, H. Pittier 5706 (Herb. N.Y. Bot. Gard.; U.S. Nat. Herb. [ 100 ] 715996); Canal Zone, Balboa. ‘*‘Never flowers a full spray. It is necessary to pick a flower at a time as they show.”’ [rec’d] May 4, 1923, C. W. Powell 67 (Herb. Ames 23965, 23966, 69927; Herb. Ames Ale. Coll. 438; Herb. Mo. Bot. Gard. 955922); Canal Zone, Fort Sherman, January 15, 1924, P. C. Standley 31231 (U.S. Nat. Herb. 1225409); Provincia de Panama, between Matias Hernandez and Juan Diaz, January 21, 1924, P. C. Standley 31944 (U.S. Nat. Herb. 1225418); Provincia de Panama, swamp between El] Jagua Hunting Club on Rio Jagua and El Congor Hill, alt. 2 m., February 10, 1935, A. A. Hunter & P. H. Allen 473 (Herb. Ames 42248); Pearl Islands, Trapiche Island, March 15, 1937, G. 8. Miller 1908 (U.S. Nat. Herb. 1688731); Pearl Islands, San José Island, March 16, 1937, G. S. Miller 1909 (U.S. Nat. Herb. 1688746) Isla Colon, April 1, 1940, HH. von Wedel s.n. (Herb. Mo. Bot. Gard. 1227010); Provincia de Bocas del Toro, vicinity of Chiriqui Lagoon, Old Bank Island, ‘flowers purplish,’? February 15, 1941, H. von Wedel 2100 (Herb. Ames 61530; U.S. Nat. Herb. 1863094); Perlas Archipelago, Gulf of Panama, San José Island (mouth of Mata Puerco), about 55 miles southeast of Balboa, April 12, 1945, J. M. Johnston 703 (Herb. Ames 64953). Trintpap: Cult. Hort. Trin. from ‘‘Inland Districts’? J. H. H[art] s.n. (Herb. Trin. 5512); Inland Woods, 1896, J. H. Hart] s.n. (Herb. Trin. 5983; Herb. Kew; Herb. Ames 66910, 68215); Brad- ford s.n. [Herb. Hance 5334] (Tyrer of EF. indivisum, Herb. Kew); Cult. Hort. Kew from Trinidad, May 1889 (Herb. Kew); Government House Grounds, June 3, 1907, W.E. Broadway s.n. (Herb. Ames 10736); Government House Grounds. “‘Flowers white,’’ June 22, 1907, W. E. Broadway s.n. (Herb. Ames 10727); Erin, March 27, 1908, W. E. Broadway s.n. (Herb. Ames 10640); Santa Cruz, Feb- ruary 23, 1912, W.E. Broadway s.n. (Herb. Kew; Herb. Mo. Bot. Gard. 918485); St. Augustine, Imperial College of Tropical Agricul- ture, April 15, 1949, N. W. Simmonds 351 (Herb. Trin. 14438; Herb. Ames 66981); St. Augustine, 6 miles from sea, W. G. Downs & T. H. G. Aitken 156 (Herb. Ames Ale. Coll, 3205a); Caigual, about 4 miles from coast, January 24, 1955, W.G. Downs & T. H. G. Aitken 15e (Herb. Ames 67831, 67783); Plain Road, 3-4 miles from sea, January 14, 1957, W. G. Downs & T. H. G. Aitken 15d (Herb. Ames Ale. Coll. 3203a). Venezurta : Near Colonia Tovar, 1856-1857, A. Fendler 2436 (Herb. Rehb. 891a; Herb. Kew; Herb. Gray 4077); [Drawing of flowering specimen and floral diagnosis] Cult. Hort. K. W. John, flowered June 1904, from Venezuela (Typr of D. venezuelanum, Herb. Ames 69928); Between Valencia and Maracay, January 31, 1918, H. Pittier 7748 (U.S. Nat. Herb. 987846); Rastrojos, near Cabudare, Lara, Decem- [ 101 |] ber 1923, J. Saer 116 (U.S. Nat. Herb. 1193211); June 1939, V. Barnes 5923 (Herb. Ames 58216). ExcLUDED OR UNCERTAIN CONCEPTS Diacrium bidentatum (Lind/.) Hemsley in God- man & Salvin Biol. Centr.-Am., Bot. 8 (1888) 221. Hpidendrum bidentatum Lindley Gen. and Sp. Orch. Pl. (1831) 98. The type of Mpidendrum bidentatum is preserved in the British Museum. Study of a photograph of the type and diagnostic sketches in the Ames Herbarium indi- cates that this concept cannot be referred to Caularthron, but that, without a doubt, it represents, as has previ- ously been suggested (Williams, L. O. in Ceiba 2 (1951) 174), Hpidendrum Boothianum dl. Diacrium Ulmckei Arinz/in Mitteil. Inst. Allg. Bot. Hamb. 6 (1927) 419. This concept was described on the basis of material which flowered in the Hamburg Botanical Garden. The plant had presumably been collected in Guatemala. Since the Hamburg Herbarium has apparently disappeared, we are unable to trace a type, if indeed an herbarium specimen were ever preserved there when the concept was described. After an examination of the description, I rather doubt that the concept can be accommodated in the genus Caularthron. [ 102 ] STUDIES IN AMERICAN ORCHIDS Il’ A NEW GENUS FROM THE COLOMBIAN AMAZONIA BY Lestige A. Garay? TIME and again, the interpretation of orchid genera has caused grave problems, since we often find that many of the species which are assigned to a particular genus are either atypical or obviously do not belong to the concept in question. Recently, I encountered a problem of this nature, which I wish to discuss here. In 1934, Mansteld described Hybochilus Huebnert, which he based on a collection sent in by Mr. Huebner from the Amazon region of Colombia. At first, it ap- peared to him that this collection might represent a new generic concept, but for some unknown reason he finally decided to include it inthe genus Hybochilus. This spe- cies has subsequently been all but unknown, and its nomenclatorial type was destroyed during the Second World War. While studying a number of collections from Colom- bia in the Ames Herbarium, I found two undetermined specimens which appeared to be quite peculiar in their 1 Previous numbers of this series were published in the Canadian Journal of Botany, vol. 34: (1956), pp. 241-260, 721-748. ? Assistant Curator of the Herbarium, Department of Botany, Uni- versity of Toronto, Toronto, Canada; at present, a Guggenheim Fel- low at the Orchid Herbarium of Oakes Ames, Harvard University. [ 103 | general habit and which immediately suggested Queket- tia, another genus of the subtribe Capanemieae and known only from the Guianas. I consequently investi- gated all of the genera and species of the Capanemieae, and I am now convinced of the identity of my material with that of the concept Hybochilus Huebneri. According to Schlechter’s system, such features as an incumbent anther and auricles which are a continuation of the clinandrium characterize the Capanemieae. A com- pletely dorsal anther and the presence of auricles on the column which enfold the stigmatic cavity are characters of the monogeneric subtribe Papperitzieae; these char- acters are clearly observable in Hybochilus Huebneri. The structure of the column, however, is much more complicated than in Papperitzia. There are two pairs of lobes: one pair is located in front of the column, with their parallel sides enclosing the stigma, the lower mar- gins being completely connate to form a shallow, cup- like cavity immediately beneath the stigmatic surface; the other pair of lobes terminate the column. At first, these are parallel and enclose the lower portion of the anther. When the anther is removed they open up slight- ly and become spirally twisted. It is hard to state the real function of these lobes, since we have no field obser- vations. It appears, however, that so long as the flowers are not visited by pollinating insects they remain fairly connivent, but after the removal of pollinia they open up slightly. I believe that these auricles protect the vis- cosity of the stigma and pollen-gland from rapid desic- cation. In view of these dissimilarities, which I consider to be of basic importance in the differentiation of subtribes as well as of genera within the subtribes, I believe that Hy- bochilus Huebneri represents a distinct genus which should be placed in the Papperitzieae. | 104 | Polyotidium Garay gen. nov.’ Sepalum posticum asepalis lateralibus omnino liberum, valde concavum; sepala lateralia usque ad medium con- nata, basi leviter saccata. Petala plana, sepalo postico subsimilia. Labellum basi columnae adnatum, non articu- latum; hypochilum late excavatum, epichilum cuneato- flabellatum, planum. Columna apoda, brevis, paulo arcu- ata, apicem versus leviter dilatata, alis quaternis ornata; alae duae inferiores in facie columnae juxta orificium stigmaticum patentes, superiores duae in apice columnae erectae. Orificium stigmae ellipticum, valde concavum. Rostellum valde conspicuum, alte bifidum. Clinandrium posticum, vix rite evolutum, integerrimum. 0.1-0.15 mm. apicem versus leviter expansus: stigma truncatum: cap- sulum. bivalve: semina ellipsoidea 0.40.2-0.25 mi. laxe elongato-areolata. Siphanthera Hostmannu (synonymous with iS. Tate: Gleason and WS. capitata Gleason) and its poorly distin- guished relatives GS. tenera Pohl, S. subtilis Pohl, JS. pratensis Met., S. ramosissima Cogn, ex Hoehne, 8. par- aensis Huber) all have varying amounts of gland-tipped trichomes. The descriptions and illustrations of all ex- cept Siphanthera Hostmannu, S. paraensis and iS. ramo- sissima omit mention of staminodia. All relatives of JS. vaupesana, except S. pratensis and S. subtilis, have wide- pored and essentially erostrate anthers. CotombBia: Comisaria del Vaupés, Rio Vaupés, Raudal de Yurupari. On sand or quartzite base. November, 1951. Richard Evans Schultes & Isidoro Cabrera 19723 (Tyrer in N.Y. Bot. Gard. Dupricate type in U.S. Nat. Herb. No. 2,195,539). My RSINACEAE Conomorpha lithophyta FP. 1. Schultes sp. nov. Arbuscula saxicola parva, usque ad decem ped. alta, pauce ramosa, ramulis junioribus minute albido-tomen- tellis, mox glabrescentibus, ramis cortice cinereo cum [ 166 ] petiolorum cicatricibus conspicuis obtecto. Folia vivo aliquid reclinata; petiolis fuscis, sulcatis, minutissime tomentellis, 2-8 cm. longis, plusminusve 8-4 mm. in diametro; lamina valdissime coriacea, obovata, conspicue marginata, apice obtusa vel rotundata, statu adulto 9-13 em. longa, 5-6.5 em. lata, supra vivo atroviridia (sicci- tate stramineo-fusea), nitida glabraque, nervis haud con- spicuis, subtus pallidiora, apparenter glabra sed per vit- ream minutissime et remote albido-lepidota, nervo cen- trale valde elevato. Inflorescentiae strictae, saepe rigide arcuatae, numerosae, foliis breviores, densissime multi- florae, usque ad basim florigerae, axi aliquid crassiusculo, dense et molliter aureo-tomentello. Flores sessiles, pa- tentes, plerumque 4 mm. longi, ore 4—4.5 mm. in diam- etro, albo-lutel, fragrantissinu. Sepala base connata, lobis triangularibus, acutis, plusminusve 1.8 mm. longis, 1 mim. latis, extus dense aureo-glanduloso-tomentellis, in- tus minute pulverulentis. Petala media pro parte con- nata, lobis crassioribus, ovatis, apice obtusis, 4 mm. longis, 1.8-2 mm. latus, extus sed intus densiore albido- glanduloso-papillosis. Stamina petalis breviora; filamen- tis crassioribus et petala adnatis; antheris recurvis, 0.6 mm. longis. Ovarium lagenitorme, 0.7-0.8 mm. in di- ametro, bene lepidatum, stylo cylindrico, crasso, usque ad 1.5 mm. longo. Fructus adhuc ignotus. The specific epithet of Conomorpha lithophyta ( dwelling plant™’) refers to the habitat chosen by this small, bushy treelet. The Savannah of Yapoboda has a base of metamorphosed quartzite of probable Cretaceous age. The endemic flora which covers it is highly adapted a xerophytism due primarily to the stone- to xerophytism lack of soil and consequent mechanical loss of rain-water. Conomorpha lthophyta inhabits the most inhospitable parts of the savannah and exhibits a predilection for rock outcroppings (see Plate NN NII, lower figure). + | [ 167 The closest ally of Conomorpha lthophyta seems to be C. magnoliifola Mez, a species of British Guiana and Surinam. The former can be distinguished from the latter at once by its very compact and densely many-flowered spikes (in contrast to loosely fewer-flowered racemes). There are also minor floral differences, such as the tri- angular lobes of the calyx and the lageniform ovary in Conomorpha lithophyta (in contrast to ovate lobes and a subglobose ovary in C. magnoliufolia). Like Conomorpha magnolufola in Surinam, this new species is reported to be employed as a minor fish-poison. The Kubeos living along the uppermost course of the Rio Kuduyari sometimes crush the leaves of Conomorpha lithophyta for use in fishing in stagnant lagoons. ‘The Kubeo name of the tree is dd-pee-ha-hee, meaning savan- nah (dd) fish-poison (pee-ha-hee). CotomBia: Comisaria del Vaupés, Rio Kuduyari, Savannah of Yap- oboda. *"Tree 10 feet tall. Flowers fragrant, yellow. Inflorescence frequented by bees.”*’ September 4, 1956. Arthur F. Barclay & Rich- ard Evans Schultes 560 (Tyre in Herb. Gray ).—Same locality and date, Barclay & Schultes 543,—Same locality and date, Barclay & Schultes 566. APOCYNACEAE Dr. Robert Woodson has contributed the descriptions and discussions of the apocynaceous plants which are dis- cussed in the following lines. Mandevilla annulariifolia JWoodson sp. nov. Frutex volubilis; ramis crassis vel crassiusculis Juven- tate minute papillatis mox cortice valde anguloso satur- ate rubro-vernicoso tectis, ramulis saepe verticillatis. Folia verticillata saepissime quaternata subsessilia line- aria 2-5 em. longa, 1-4 mim. lata, firmiter membranacea glabra nervo medio supra distante glanduligero, Inflores- centia terminalis vel subterminalis pauciflora: pedunculo [ 168 | vix 1 em. longo minute puberulo; pedicellis ca. 2 mm. longis minute puberulis: bracteis minute ovatis vix 1 mim. longis. Calycis laciniae ovatae acutae ca. 1.5 mm. longae minute papillatae intus basi squamella profunde lacerata munitae. Corolla infundibuliformis lutea extus minutissime papillata: tubo proprio ca. 2 cm. longo basi ca. 1 mm. diam. paululo gibboso; faucibus tubulo- ‘ampanuliformibus ca. 1.7 cm. longis, ostio ca. 0.6 em. diam. : lobis late dolabriformibus patulis ca. 1.2 > ¢m. longis. Antherae late oblongo-sagittiformes basi rotun- date 2-lobae ca. 3.5 mm. longae apice minutissime bar- batae caeterumque glabrae. Ovaril carpella vix 1 mm. longa glabra nectario late 5-lobo ovario ca. dimidio aequante; stigmate pentagono-umbraculiformi obscure apiculato ca. 1 mm. longo. Folliculi gracillimi faleati obscure articulati usque 10 em. longi. Mandevilla annularifolia is closely allied, indubitably, to M. Benthamiu of the Roraima shield of interior Vene- zuela and Guiana, which are low erect shrubs with some- what broader and thicker leaves and more elongate in- florescences. The suggestively antediluvian appearance of Mandevilla annularifolia is imparted by the narrow, whorled leaves borne in dense tufts at the ends of the thick, frequently whorled branchlets. Cotompia: Comisaria del Vaupés, Rio Apaporis, Cachivera de Jiri- jirimo y alrededores. Altitude about 250 m. September 16, 1951. Richard Evans Schultes & Isidoro Cabrera 14013 (Tyrer in Mo. Bot. Gard.).—Rio Apaporis, Raudal de Jirijirimo, November 27, 1951. Schultes & Cabrera 146.27.—Rio Apaporis, Raudal de Jirijirimo, below mouth of Rio Kananari, December 8, 1951. Schultes & Cabrera 1050.3, —Comisaria del Vaupés, Rio Piraparana, tributary of Rio Apaporis, September 18, 1952. Schultes & Cabrera 17536, Mandevilla cuneifolia Woodson sp. nov. Frutex volubilis: ramulis gracilibus glabris. Folia decussata breviter petiolata; lamina obovata vel elliptico- obovata abrupte acuminata basi acuta 4-7 cm. longa, (169 | EXPLANATION OF THE ILLUSTRATION Phare XXXI, Conomorpua tirnopuyta FP. F. Schultes. 1, flowering branch, slightly less than one half natural size. 2, portion of inflorescence, showing two flowers, approximately two and one half times natural size. 38, petals opened, approxi- mately twice natural size. Drawn by Ruru Barron [ 170 ] PLatE XXXI CONOMORPHA 7B lithophyla RE Rhutlles EXPLANATION OF THE ILLUSTRATION Prare XXXII. (Upper) View of the quartzitic sa- vannah at Yapoboda, upper Rio Kuduyari, Com- isaria del Vaupés, Colombia—type locality of Conomorpha lithophyta. (Courtesy American Orchid Society, Inc. ) (Lower) Conomorpua tirnopuyta PR. &. Schultes. Flowering branch of the type plant. Photographs by Racuarp Evans Scuuirrs PLATE XXXII # EXPLANATION OF THE ILLUSTRATION Prare XXXIII. Conomorpua tirnopnytra PR. FE. Schultes. Habit photograph of the type plant. Photograph by Ricuary Evans Scuvuires PLATE Nw LET 1.52.5 em. lata firmiter membranacea glabra nervis se- condaris arcuatis sat remotis nervo medio supra pauci- glanduligero; petiolo 8-5 mim. longo. Inflorescentia lat- eralis vel subterminalis pauciflora glabra; pedunculo 1— 2.5em. longo; pedicellis subnullis; bracteis minutissime ovatis. Calycis laciniae ovato-trigonales longe acuminatae extus papillatae intus squamella deltoidea minute erosa munitae. Corolla infundibuliformis lutea extus omnino glabra: tubo proprio 8.5-4 ¢m. longo basi ca. 1.5 mm. diam. ; faucibus conicis ca. 2.5 em. diam., ostio ca. 1 em. diam. ; lobis late dolabriformibus ca. 2 ¢m. longis patulis. Antherae obovato-oblongae basi breviuscule auriculatae ‘a. 8 mm. longae dorso glabrae. Ovaril carpella ovoidea ‘a. 1.5 mm. longa, nectariis 5 plusminusve connatis subae- quilongis; stigmate pentagono-umbraculiformi obscure apiculato ca. 1 mm. longo. A member of the Mandevilla scabra alliance, M. cunei- folia is distinguished by the obovate or elliptic-obovate cuneate leaves and reduced inflorescence. Cotompia: Comisaria del Vaupés, Rio Piraparana (tributary of Rio Apaporis), Cafiio Teemeefia, Savannah O-koo-me-gwa, September 6, 1952, Richard Evans Schultes & Isidoro Cabrera 17234 (Tyrr in Mo, Bot. Gard. ).—Comisaria del Vaupés, Rio Vaupés, between Mitt and Javareté, Yutica, May 14-24, 1953. Schultes & Cabrera 19877. Mandevilla stephanotidifolia ]0odson sp. nov. Frutex volubilis; ramulis teretibus vel paulo compres- sis glabris. Folia decussata petiolata; lamina latiuscule oblongo-elliptica in acumine breve abrupte contracta basi obtusa vel rotundata 6-12 cm. longa, 4-7 em. lata sub- coriacea subglauca glabra nervo medio supra distante glanduligero; petiolo ca. 1 em. longo. Inflorescentia aut terminalis aut lateralis spicata multiflora: pedunculo ca. 6-12 em. longo crassiusculo glabro; pedicellis nullis vel vix bene manifestis: bracteis speciosis petalaceis (albis 4) caducis late oblongo-obovatis carinatis apice valde acu- [ 176 ] minatis sessilibus 2.5-8 em. longis. Calycis laciniae late ovatae acutae ca. 1 mm. longae extus minutissime pap- illatae intus basi squamella truncate deltoidea praeditae. Corolla infundibuliformis lutea extus glabra vel indis- tincte papillata tubo laeviter arcuato-gibboso; tubo pro- prio ca. 83cm. longo basi ca. 2 mm. diam.; faucibus con- icis ca. 1.5 cm. longis, ostio ca. 1 cm. diam. ; lobis late dolabriformibus ca. 1.7 em. longis patulis. Antherae oblongo-ellipticae basi obtuse auriculatae ca. 6.5 mm. longae dorso glabrae. Ovarii carpella gemina oblongo- fusiformia ca. 2 mm. longa indistincte papillata basi nec- tariis 5 subquadratis carnosis cincta; stigmate pentagono- umbraculiformi longe apiculato ca. 4 mm. longo. Folli- culi graciles falecati apice saepe coaliti obscure monili- formes 8-10 cm. longi glabri. Mandevilla stephanotidifolia is closely related to M. Steyermarkii of the Roraima shield, which, however, has slenderer stems, thinner, cordate-ovate leaves, and a slenderer inflorescence. CotompiA: Comisaria del Vaupés, Rio Apaporis, Cachivera de Jiri- jirimo y alrededores. Altitude about 250 m., September 16, 1951. Richard Evans Schultes & Isidoro Cabrera 14010 (Tyrr in Mo. Bot. Gard. ).—Rio Apaporis, Raudal Jirijirimo, below mouth of Kananari. Altitude about 900 ft. January 21, 1952. Schultes § Cabrera 14952. —Rio Apaporis, Raudal de Jirijirimo, November 27, 1951. Schultes & Cabrera 14632.—Comisaria del Amazonas, Rio Caqueta, La Pedrera and vicinity, May 2, 1952. Schultes & Cabrera 16325. Mandevilla thevetiocides Woodson sp. nov. Frutex humilis; ramulis crassis Juventate ancipitibus minute puverulo-papillatis mox cortice atrorubescenti tectis. Folia decussata brevissime petiolata lineari-ellip- tica cum apice tum basi anguste acuta 4-8 cm. longa, 5-10 mm. lata, coriacea margine revoluta venis secund- ariis multis horizontalibus supra illustria nervo medio pauciglanduligero subtus minutissime velutina; petiolo bun ca. 2mm. longo. Inflorescentia aut terminalis aut sub- terminalis pauciflora minute puberula; pedunculo crasso 1-2 em. longo; pedicellis 2-8 mm. longis; bracteis mi- nute ovatis caducis. Calycis laciniae ovatae acutae ca. 2 mm. longae extus puberulo-papillatae. Corolla infundi- buliformis alba extus papillata; tubo proprio gracillimo ca. 8 cm. longo basi ca. 1 mm. diam. ; faucibus anguste conicis ca. 1.5 em. longis, ostio ca. 7 mm. diam. ; lobis oblique dolabriformibus ca. 2 em. longis patulis. Anth- erae late oblongo-sagittatae ca. 4 mm. longae basi obtuse bilobatae dorso glabrae. Ovarii carpella oblongoidea ca. 1.5 mm. longa glabra nectario 5-lobo basi cineta; stig- mate pentagono-umbraculiformi obscure apiculato ca. 1.5 mm. longo. Closely allied to Mandevilla nerioides, this new species may be distinguished by the dense horizontal secondary venation of the leaves, minutely velutinous beneath. Cotoms1a: Comisaria del Vaupés, Rio Kananari (affluent of Rio Apaporis), Cerro Isibukuri. October 29, 1951. Richard Evans Schultes & Isidoro Cabrera 14508 (Tyre in Mo. Bot. Gard.). Prestonia vaupesana JW oodson sp. nov. Frutex volubilis omnino glaber; ramulis teretibus gracillimis. Folia decussata petiolata; lamina ovato- elliptica acuminata basi obtusa 7-12 cm. longa, 3-6 cm. lata firmiter membranacea; petiolo 5-7 mm. longo. In- florescentia simplex corymbiformis interpetiolata pluri- vel multiflora; pedunculo gracile 7-16 cm. longo; pedi- cellis gracillimis ca. 1.5 cm. longis in fructu accrescenti- bus; bracteis minimis. Calycis laciniae anguste oblongo- lanceolatae 7-8 mm. longae herbaceae apice saturiorae intus basi squamella truncata munitae. Corolla aut gilva aut livide rosea salverformis; tubo ca. 1.5 em. longo basi ca. 2.5 mm. diam., annulo conspicuo ca. 1 mm. alto minute papillato, appendicibus epistaminalibus anguste oblongis ca. 2 mm. longis ostio vix attingentibus; lobis [ 178 ] late dolabriformibus ca. 9 mm. longis refractis. Antherae dimidio exsertae lanceolatae ca. 5 mm. longae dorso sparsiuscule pilosulae. Ovaril carpella ca. 1 mm. longa; nectarlis carnosis basi connatis ovario aequantibus. Fol- liculi gracillimi faleati obscure articulati. The description of new species of Prestonia upon sus- piciously slight characters and from unicate specimens is a recurring annoyance. Although Prestonia vaupesana is very closely related to P. Lindleyana W oods., to which it will key out in my revision of the genus (in Ann. Missouri Bot. Gard. 23 (1986) 276), the several collec- tions made by Schultes and Cabrera, all from the same general area in the valley of the Rio Apaporis, give ample evidence of the characters of a natural population. From Prestonia vaupesana, P. Lindleyana differs in its some- what more coriaceous, oblong- or obovate-elliptic foliage, in its shorter inflorescence with stouter peduncles and pedicels, and particularly in its rather broadly ovate, more substantial calyx lobes which, with the pedicels, are prominently suffused with purple. Cotomsia: Comisaria del Vaupés, Rio Apaporis, Cachivera de Jiri- jirimo y alrededores. Altitude about 250 m. September 16, 1951. Richard Evans Schultes & Isidoro Cabrera 14039 (Tyrer in Mo. Bot. Gard.).—Rio Apaporis, Soratama above mouth of Rio Kananari and vicinity. Altitude about 900 ft. January 1952. Schultes & Cabrera 19611.—Rio Apaporis, Raudal Yayacopi (La Playa) and vicinity. Altitude about 800 ft. August 18, 1952. Schultes & Cabrera 1697 4.— Rio Apaporis entre el Rio Pacoa y el Rio Kananari. Altitude about 250m. July 5, 1951. Schultes & Cabrera 12893. Tabernaemontana rimulosa Woodson sp. nov. Fruticulus humilis, ramulis crassiusculis juventate glabris mox cortice tenui griseo-fulvo longitudinaliter exfoliato tectis. Folia decussata subsessilia in paribus aequalibus; lamina anguste oblongo-elliptica apice acuta basi more subpetiolato gradatim decurrente 12-13 cm. longa, 3-3.7 cm. lata coriacea margine leviter revoluta [ 179 ] omnino glabra supra illustria subtus opaca lutescentia reticulo venularum fere omnino immerso. Inflorescentia interpetiolatis subterminalis subsessilis glomerate com- posita pluriflora; pedunculis pedicellisque perbrevibus; bracteis minimis glabris. Calycis laciniae late ovales apice rotundatae plusminusve inaequales 2.5-8 mm. longae extus glabrae intus supra basim squamellas liguliformes 2-4 gerentes. Corollae albae lobis margine roseo-tinger- entibus; tubo cylindrico ca. 1.7 em. longo, basi ca. 1.5 mm. diam. medio staminigero parte dimidio superiore extus dense minuteque velutino-papillato basim versus glabro; lobis patulis valde oblongo-dolabriformibus ca. 1 cm. longis supra dense minute velutino-papillatis. Antherae sessiles angustissime lanceolatae longe acu- minatae ca. 5 mm. longae thecarum parte basali angus- tissima recta. Ovari carpella oblongo-fusiformia ca. 1.5 mm. longa glabra basi nectario dimidio longitudine cincta; stigmate late pentagono-umbraculiformi obtuse apiculato ca. 0.7 mm. longo. Fructus ignoti. ‘This new species is apparently most closely related to Tabernaemontana coriacea Link ex R. & S., but it differs significantly in the narrower leaves, the subsessile glom- erate inflorescences, the low stature and, particularly, in the thin, loosely exfoliating periderm of the older branches. [ 180 | BOTANICAL MUSEUM LEAFLETS HARVARD UNIVERSITY CamprinGr, Massacuusetts, November 20, 1958 Vor. 18, No. 5 HOW ABSURD CAN A NOMENCLATURAL PROPOSAL BE? BY LesuiEk A. Garay anp Ricuarp Evans ScHULTES Boranists who are interested in plants as living things and not as pretexts for legalistic juggling of nomencla- ture are becoming increasingly exasperated with the growing amount of attention paid in our congresses and in our journals to hair-splitting and often superficial in- terpretations of the International Code of Botanical Nomenclature, whose basic reason for existence is pre- cisely to standardize nomenclature and make such quib- bling unnecessary. We would be the first to recognize that rules and regulations are essential for the standard- ization of nomenclature, yet we rebel at the threat that the activity of other taxonomists become more and more subservient to the confusion wrought by those whose only interest in plants seems to be the legalism surround- ing the clarification of their proper naming. Taxon, the journal of the International Association for Plant Taxonomy, has recently been given over largely to articles on nomenclature by individuals and commit- tees. It is with one part of a recent article that we wish to deal in this note, our primary purpose being to point out one of the many absurdities which our congresses are being asked to consider. We refer to the ‘‘Report of the Committee for Spermatophyta. Conservation of generic [181 | names I”* in Taxon 7 (1958) 184-198. In this report, we find the following proposal for conservation of the name of an orchid genus: 1533. Bletilla Reichenb.f. (1851) vs. Jimensia (Raf.) (1888) (6-2) (Syn. Prop. Stockholm 228.) Bletilla has recently been recognized as a distinet genus with some 6 species. Rafinesque’s name has never been used. To adopt it would necessitate new combinations, Even a superficial reading of this proposal is enough to frighten and shock the average botanist into the reali- zation of the lack of seriousness and thoroughness which is unfortunately becoming more common in the work of those whose main interest apparently is to regulate, no matter on how trivial a point, the work of their colleagues who are concerned with the study of plants. Reichenbach described Bletil/a in 18538, giving the con- cept avery appropriate name suggestive of a resemblance to Bletia; the same concept, however, had been recog- nized and adequately treated thirteen years earlier by Rafinesque under the name Jimensia. Rafinesque’s treat- ment (FI. Tellur. 4 (1836) 88) is clear: 909. Jimensia R. (Jap. bot.) Petalis ovatis concavis, 2 internis, label. trifido emarg. basi callis 2 obl. medio concavo, col. filif. in- curva, stig. bifid. concavo, antheris 2 dorsalis, capsula clavata, Scaposa, fol. gladiatis, fl. spicatis—Type J. nervosa R. Limodorum striatum Th. fl. jap. scapo angulato, fol. rad. glad. nervosis, fl. cer- nuis, bracteatis. Japan, fl. yellow. The G. Limodorum contained many anomalies also, sp. with or without spurs, beards or no beards, many kinds of pillars or styles or clinandres &c. In 1950, before the Stockholm Congress, a Japanese botanist proposed the conservation of Bletilla. This pro- posal was not acted upon in Stockholm. It did not ap- pear in the Paris proposals. Now, the name has been resurrected and proposed for conservation once again, with the most amazing reason: ‘‘Rafinesque’s name has never been used. ‘To adopt it would necessitate new com- [ 182 | binations.”’ Is it conceivable that a committee of bota- nists would suggest deviation from the basic rule of pri- ority because an earlier name had not been used or because several new combinations might result? Naturally, most conservative botanists try to resist altering nomenclature in the cases of very large genera or in generic names which have been long or widely used in economic botany or hor- ticulture, but the genus under question is neither large nor of economic or horticultural importance. Reichenbach’s concept included two different entities. He listed two species in the genus, Bletilla florida from the New World and B. gebinae from the Old World. It is now recognized that Bletilla florida belongs in the genus Bletia. Rafinesque’s concept is concerned only with the true Bletilla element, an additional cause for taking up his name for the concept, which would have been obvious if those responsible for this proposal had any knowledge of the plants involved. In order to forestall absurd and unnecessary legislative action, we herewith make the appropriate transfers to the genus Jimensia: Jimensia formosana (Hayata) Garay & R. E. Schultes comb. nov. Bletia formosana Hayata Mat. Fl. Formos. (1911) 823. Bletilla formosana (Hayata) Schltr. in Fedde Rep. 10 (1911) 256. Jimensia kotoensis (Hayata) Garay & R. E. Schultes comb. nov. Bletia kotoensis Hayata Mat. Fl. Formos. (1911) 325. Bletilla kotoensis (Hayata) Schltr. in Fedde Rep. 10 (1911) 256. Jimensia morrisonicola (Hayata) Garay & R.E. Schultes comb. nov. [ 183 ] Bletia morrisonicola Hayata Mat. FI]. Formos. (1911) B24. Bletilla morrisonicola (Hayata) Schltr. in Fedde Rep. 10 (1911) 256. Jimensia ochracea (Sch/tr.) Garay & R. 1. Schultes comb, nov. Bletilla ochracea Schltr. in Fedde Rep. 12 (1918) 105. Jimensia scopulorum (W.W Sm.) Garay & RL. Schultes comb. nov. Pleione scopulorum W.W.Sm. in Notes R. Bot. Gard. Edinb. (1921) 218. Bletilla scopulorum (W. W. Sm.) Schltr. in Fedde Rep. 19 (1924) 875. Jimensia sinensis (Rolfe) Garay & R.E. Schultes comb, nov. Arethusa sinensis Rolfe in Journ. Linn. Soc. Bot. 36 (1903) 46. Bletilla sinensis (Rolfe) Schltr. in Fedde Rep. 10 (1911) 256. Bletilla chinensis Schltr. Die Orchideen (1914) 107 sphalm. Jimensia striata (Thunb.) Garay & R.E.Schultes comb, nov. Limodorum striatum Thunb. FI. Jap. (1784) 28. Epidendrum tuberosum Lour. Fl. Cochin. (1790) 528. Epidendrum striatum (Thunb.) Thunb. in Trans. Linn. Soc. pt. 2 (1790) 327. Cymbidium striatum (Thunb.) Sw. in Nov. Act. Upsal. 6 (1797) 77. Cymbidium hyacinthinum Sm. Exot. Bot. 1 (1804) 117. Gyas humilis Salisb. in Trans. Hort. Soc. 1 (1812) 300. [ 184 ] Bletia hyacinthina (Sm.) R. Br. in Ait. Hort. Kew., ed. 2, 5 (1813) 206. Jimensia nervosa Raf. Fl. Tellur. 4 (1836) 88. Bletia gebina Lindl. in Journ. Hort. Soe. 2 (1847) 307. Bletilla gebinae (Lind].) Rehb.f. in Fl. Serres, ser. 1, 8 (1853) 246, Calanthe gebinae (Lindl.) Lodd. ex Rehb.f. in FI. Serres, ser. 1, 8 (1858) 246. Bletilla striata (Thunb. ) Rehb.f. in Bot. Zeit. 86 (1878) 795. Jimensia szetschuanica (Sch/tr.) Garay & R. E. Schultes comb. nov. Bletilla szetschuanica Schltr. in Fedde Rep. Beih. 12 (1922) 844. Jimensia yunnanensis (Schltr.) Garay & R. FE. Schultes comb. nov. Bletilla yunnanensis Schltr. in Fedde Rep. Beih. 12 (1922) 843. Bletilla yunnanensis (Schltr.) var. Limprichtu Schltr. in Fedde Rep. Beih. 12 (1922) 344. EXCLUDED SPECIES Bletilla florida Rchb.f. in FI. Serres, ser. 1, 8 (1853) 246= Bletia florida R.Br. Bletilla japonica (A.Gray) Schltr. in Fedde Rep. 10 (1911) 256= Arethusa japonica A. Gray. [ 185 ] STUDIES IN AMERICAN ORCHIDS IV.' BY LEsLIE A. GARAY In the following pages, various new species are described and several new combinations of taxonomic importance are proposed, all of which were discovered during routine checking of collections from the Andes. The arrange- ment of genera follows the phylogenetic system proposed by Rudolf Schlechter in Notizblat des Botanischer Gar- tens und Museum zu Berlin-Dahlem 9 (1926) 563-591. Duckeella pauciflora Garay sp. nov. Saxicola, usque ad 50 cm. alta; radicibus satis crassi- usculis, leviter puberulis; foliis 2-8 basilaribus, lineari- lanceolatis, satis coriaceis, 15-80 cm. longis, 8-5 mm. latis; scapo simplici vel in parte superiore pauciramoso, erecto, stricto, 830-50 cm. alto; inflorescentia abbreviata, pauciflora; bracteis navicularibus, lanceolato-triangulari- bus, ovariis pedicellatis multoties brevioribus, 4-5 mm. longis; floribus satis magnis, succedaneis, flavis; sepalo postico oblongo-obovato, concavo, reflexo, apice acuto, 7-nervio, 20-22 mm. longo, 6-7 mm. lato; sepalis lateral- ibus leviter obliquis, oblongo-obovatis, concavis, reflexis, 7-nerviis, 20-22 mm. longis, 6-7 mm. latis; petalis valde 1 Previous numbers of this series were published in the Canadian Journal of Botany, vol. 34 (1956) 241-260, 721-748 and in Bot. Mus. Leafl. Harvard Univ. 18 (1958) 103-108. [ 186 ] tenuis, late ovato-rhombeis, acutis vel obtusis, reflexis, 9-nerviis, 18-21 mm. longis, 8-12 mm. latis; Jabello complicato, 3-lobo; lobis lateralibus oblongis, obtusis vel subacutis, 3.5-4 mm. longis, lobo intermedio ovato- oblongo, acuto vel obtusiusculo, 14-17 mm. longo, 5-8 mm. lato; disco supra basim multi-cristato, cristis antice in fimbrias dissolutis; columna 8 mm. longa, apice bi- auriculata; ovariis pedicellatis 1.5-2 em. longis. This is the second species of the genus to be noted. A third, as yet undescribed, from the Guiana Highlands, is under study at present by Mr. Schweinfurth. Duckeella pauciflora differs from D. Adolfii Campos Porto & Brade in having a short raceme, larger flowers and differently shaped petals and lip. CoLtompia: Comisaria del Vaupés, Rio Kubiydé (tributary of Rio Vaupés); Cerro Kafiendaé, savannahs about 15 miles upstream from mouth, Quartzite base. Altitude about 800-900 feet. Nov. 10, 1952. Richard Evans Schultes & Isidoro Cabrera 18309 (Tyr in Herb. Ames no. 69675; Dupticare Type in U.S. Nat. Herb.); Schultes & Cabrera 18399 (Herb. Ames no. 69674).—Yapobodaé. P. H. Allen 3203 (U.S. Nat. Herb. no. 1951970). Sobralia lancea Garay sp. nov. Saxicola, usque ad 50 cm. alta; radicibus fasciculatis, satis crassis pubescentibus; caulibus caespitosis, erectis, parte basali vaginatis, supra paucifoliatis, teretibus, gla- bris; foliis lanceolatis, apice longe acuminatis, basi in vaginam caulem amplectentem angustatis, subcoriaceis, plicatis, 16-25 em. longis, 1-1.5 cm. latis; inflorescentia valde abbreviata, bracteis lanceolatis, bifariis, arctissime adpressis omnino obtecta; floribus succedaneis, tenuis- simis, glabris; sepalis petalisque inter se simillimis, oblongo-lanceolatis, brevissime acuminatis, 5-7 nerviis, 3 cm. longis, 5 mm. latis; labello ovato-pandurato, basi rotundato, apice undulato; disco a basi usque ad con- strictionem bilamellato, lamellis leviter crenulatis, supra dentiformibus, 2.4 em. longo, supra basim 12 mm., ad [ 187 ] EXPLANATION OF THE ILLUSTRATION Prare XXXIV. Duckerecia paucirtora Garay. 1, plant, one third natural size. 2, flower, expanded, natural size. 3, lip, twice natural size. Drawn by Rutu Barron [ 188 ] PuatTE XXXIV DUCKEELLA pauciflora Garay JB apicem 8 mm. lato; columna gracili, alata, 1 em. longa, faleculis nullis; ovariis graciliter pedicellatis, usque ad 2.5 em. longis. Apparently, the Peruvian Sobralia candida (Poepp. & Endl.) Rehb.f. is the only species in the genus with which this new taxon might be confused. Vegetatively these two species are very similar, though iS. /ancea pos- sesses longer and more acuminate leaves. In the floral structures, however, a number of discrepancies can be observed: the sepals and petals are acute in S. candida and acuminate in SS. /ancea; in the former the lip is 3- lobed, with the apical lobe wider than the lateral ones; in the latter the lip is pandurate, and the apical portion is much narrower than the basal part. In addition to these differences, S. candida is described as having ‘‘snow- white’? flowers, while the color of S. /ancea is canary- yellow with purple lamellae on the lip. Cotompra: Auchicaya (Cali-Buenaventura), 500 meters alt. August 11, 1952. M. Kéie 4787 (Tyrr in Herb. Ames no, 68437; Dupiicare tyre in Herb, Bot. Mus. Univ. Copenhagen), Erythrodes cylindrostachys Garay sp. nov. Terrestris, usque ad 80 cm. alta; rhizomate longe re- penti, satis carnoso, crasso; radicibus crassiusculis, levi- ter flexuosis, puberulis, albidis; caulibus suberectis vel ascendentibus, primum remote vaginatis, deinde 8—5- foliatis, glabris; foliis ovato-lanceolatis, acuminatis, basi in petiolum attenuatis, mox in vaginam caulem amplec- tentem dilatatis, 14-20 cm. longis, 2—4.5 em. latis; in- florescentia cum pedunculo ca. 2 em. longo densissime puberula, valde compacta, cylindracea, 9-13 cm. longa; bracteis lanceolatis, acuminatis, ovariis brevioribus, 1.6 em. longis; floribus satis magnis, albidis; sepalo postico ovato-lanceolato, acuto, concavo, 8-nervio, intus glabro, extus sparse glanduloso-hirsuto, 1.2 cm. longo, 4 mm. lato; sepalis lateralibus reflexis, leviter obliquis, ovato- [ 190 ] lanceolatis, 3-nerviis, intus glabris, extus glanduloso- hirsutis, 1.2 cm. longis, 4 mm. latis; petalis ovato- spathulatis, apice acutis vel obtusis, uninerviis, intus extusque glabris, cum sepalo postico galeam formanti- bus, 1.2 em. longis, 8 mm. latis; labello trilobo, lobis lateralibus quadratis, antice obtusis, lobo intermedio re- curvo, quadrato-reniformi, brevissime apiculato, margine ais > sy LARS CAK TENS. ERYTHRODES CYLINDROSTACHYS Garay valde undulato, 9 mm. longo, antice 7 mm. lato; caleare eylindrico, leviter sinuato, acuto, 38 cm. longo; ovario cylindrico, glanduloso-hirsuto, 2.2 cm. longo. This new species has no close alliance among the known taxa. The shape of the lip, together with the long and thick spur, is very different from that in any concept yet described. Cotomsia: Departamento del Valle, Cordillera Occidental, vertiente occidental; monte La Guarida, filo de la cordillera sobre La Carbonera (entre Las Brisas y Alban). 1950-2000 meters alt. Oct. 16, 1946. J. Cuatrecasas 22232 (Typr in Herb. Ames).—La Cumbre. 1800-2100 meters alt. May 14, 1922. F. W. Pennell & E. P. Killip 5769 (U.S. Nat. Herb. no. 1141933; Herb. Ames no. 47353). [ 191 ] Erythrodes quadrata Garay sp. nov. Terrestris, usque ad 50 cm. alta; radicibus satis cras- sis, villosulis; caulibus leviter ascendentibus vel erectis, dimidio inferiori 5—G6-foliatis, supra remote vaginatis; foliis erecto-patentibus, petiolatis; lamina ovata vel ovato-lanceolata, acuta, basi abrupte vel sensim in petio- lum angustata, 7-10 cm. longa, 2.5-4 cm. lata; racemo ERYTHRODES QUADRATA Garay cylindrico, satis dense multifloro, ca. 8 cm. longo; brac- teis lanceolatis, acuminatis, ovariis aequilongis seu paulo brevioribus; floribus in genere inter minores; sepalo pos- tico ovato, obtuso, uninervio, concavo, intus glabro, extus sparse piloso, 83 mm. longo, 1.5 mm. lato; sepalis later- alibus faleato-oblongis, acutis vel obtusis, uninerviis, intus glabris, extus sparse pilosis, 8.5 mm. longis, 1.5 mm. latis; petalis cum sepalo postico galeam formantibus, obovatis, acutis, vel obtusis, basim versus attenuatis, glabris, 3 mm. longis, 0.75 mm. latis; labello quadrato- oblongo, basi valde concavo, supra medium utrinque plicato, antice truncato et in medio brevissime apiculato, 3mm. longo, antice 1.5 mm. lato; disco a basi usque ad [ 192 | plicam unicarinato; calcare porrecto, clavato, 2 mm. longo, 1 mm. crasso, ovario tertia parte breviore. This new species may be compared with the Colom- bian Hrythrodes zeuxinoides (Schitr.) Ames, but differs from it in having much smaller flowers and a very dis- similar lip. The flowers, judged from the specimen, are greenish-white in color. Cotomspra: Departamento del Magdalena, Cordillera Oriental, Sierra de Perija, 6 km. east-northeast of Manaure, 42 km. east of Valledu- par, 7 km. from the Venezuelan border. 2100 meters alt. Feb. 1, 1945. Martin L. Grant 10748 (Type in U.S. Nat. Herb. no. 2107290). Ecuapor: Chimborazo, Riv. Chasuan. R. Spruce 6138. Herb. Reich- enbach nos. 37766, 37767. Erythrodes xystophylloides Garay sp. nov. Terrestris, erecta, 20-25 cm. alta; rhizomate cauli- formi, repenti; radicibus ex internodiis orientibus, fili- formibus, leviter flexuosis, sparse pubescentibus; caull- bus erectis, primum remote vaginatis, deinde 4-6-foliatis ; foliis anguste lanceolatis, subacutis vel acuminatis, basi attenuatis, deinde in vaginas dilatatis transeuntibus ; ERYTHRODES XYSTOPHYLLOIDES Garay [ 193 ] lamina 2.5-8 cm. longa, 4-5 mm. lata; pedunculo erecto, remote 8-vaginato; racemo pauci- (6-8-) floro; bracteis lanceolato-setaceis, ovariorum pedicellatorum dimidium aequantibus; floribus in genere inter minores, albidis; sepalo postico ovato, obtuso, valde concavo, uninervio, extus sparse pilosulo, 5 mm. longo, 1.75 mm. lato; sepalis lateralibus ovato-oblongis, leviter obliquis, obtusis, extus sparse pilosis, uninerviis, 5 mm. longis, 1.5 mm. latis; petalis oblique spathulatis, uninerviis, glabris, quarta parte apicali sese tegentibus et conglutinatis, 5 mm. lon- gis, 1 mm. latis; labello conduplicato, e basi elliptico- cucullato apice in laminam anchoriformem rotundatam subito dilatato, 5 mm. longo, in medio 2 mm., apice 4 mm. lato; calcare gracili, 4 mm. longo, 1 mm. crasso. In general appearance this new species seems to be a miniature form of Hrythrodes aystophylla (Rchb.f. ) Ames. These two species may be distinguished from each other, apart from size, by the shape of the lip, which is bilobed in EL. wystophylla and anchoriform in the new concept. In natural position, the lip is conduplicate- cucullate with the anchoriform tip reflexed and rolled inwardly. CotomBiA: Dept. Cundinamarca; Cordillera Oriental, Ojo de Agua, south side Guavio River, 32 km. east of Gachalé. 2035 meters alt. Nov. 1, 1944. Martin L. Grant 10557 (Tyrer in Herb. Ames no. 69102; Dupticare tyre in U.S. Nat. Herb.). Stelis nexipous Garay sp. nov. Epiphytica, caespitosa, usque ad 25 cm. alta; radicibus filiformibus flexuosis, glabris; caulibus secundariis erectis vel paulo arcuatis, vaginis 8 alte amplectentibus maxi- ma pro parte obtectis, 6-12 cm. longis; folio coriaceo, oblongo-ligulato, apice obtusiusculo, basi in petiolum brevem sensim angustato, petiolo incluso 6-10 cm. longo, 0.7-1.2 cm. lato; inflorescentia singula, erecta vel inter- dum paulo arcuata, usque supra basim multiflora, 8-14 [ 194 | em. longa; bracteis infundibuliformibus, acutiusculis, quam ovaria pedicellata longioribus, 8-4 mm. longis; sepalis lateralibus inter se liberrimis sed cum sepalo pos- tico maxima pro parte in synsepalum connatis; synsepalo antice 3-lobo, per lobum 8-nervio, intus minutissime hir- sutulo, ad basim leviter cochleato, 1-1.3 cm. longo, inter lobis laterales 1 cm. lato; petalis valde carnosis, cuneato- flabellatis, obtusis, dimidio superiore valde incrassatis, basi 8-nerviis, 0.75 mm. longis, 1.25 mm. latis; labello valde carnoso, triangulari, basi valde incrassato, ante in- crassationem leviter excavato, 0.75 mm. longo latoque; columna humili, crassa, sursum abrupte dilatata, brachiis divaricatis; ovario cum pedicello, 2-8 mm. longo. Stelis nexipous is rather peculiar in the shape of its sepals. In appearance it does not resemble any other species in the genus. The genus Physothallis, which was established on a similar monosepalous character, has a column typical of the genus Physosiphon, and I am now convinced that the two genera are inseparable. How- ever, the concept Physothallis is just as abnormal in the genus Physosiphon as the species described herein is in the genus Stelis. A few more cases like that of S. neax- pous will help to confirm my feelings that we are dealing in the Pleurothallidinae with a very natural biological unit or genus which encompasses all the generic concepts proposed up-to-date within that subtribe. Ecuapor: Prov. Napo-Pastaza, Mera, towards Mangayacu; epi- phyte in rastrojo. Ca. 1100 m. alt. March 7, 1956. Erik Asplund 19638 (Tyrer in Herb. State Mus. Nat. Hist., Stockholm, Sweden; Dupti- CATE TYPE in Herb. Ames). Platystele ornata Garay sp. nov. Epiphytica, caespitosa, pusilla, usque ad 2.5 cm. alta; radicibus filiformibus flexuosis, glabris; caulibus secun- dariis vix rite evolutis, ullis, vagina unica omnino obtec- tis; folio obovato-spathulato, apice obtuso, ca. 2 cm. [ 195 | EXPLANATION OF THE ILLUSTRATION Pirate XXXV. Srevis nexipous Garay. 1, plant, one half natural size. 2, flower, three times natu- ral size. 8, petal, fifteen times natural size. 4, lip, twenty times natural size. Drawn by Evmer W. Suiru [ 196 ] PuareE XNNXV STELIS MEXUPOUS PLaTE XXXVI ame Anther & Pollinia Column mm Copsule Vm Apex of Leaf WN4 rev PLATYSTELE ORNATA Garay Drawn by G. C.K. Dunsrervitie longo, 3 mm. lato; inflorescentia capillari, erecta vel paulo arcuata, subdense pluriflora, ca. 2 cm. longa; bracteis infundibuliformibus, 0.1 mm. longis; sepalis ovatis, acutis, extus sparse glanduloso-pilosulis, ca. 0.5 mm. longis; petalis obovatis, apice acutis, margine glanduloso-pilosulis, sepalis paulo brevioribus; labello ovato-lanceolato, leviter cochleato, apice acuminato, sepalis magnitudine, disco nudo; columna humili, crassa, generis; ovario cum pedicello quam bracteis longiore. This new species is easily separable from the others in the genus by its glandular-hirsute floral segments. VENEZUELA: New road, Santa Teresa to Altagracia. Dec. 28, 1953. G. C. K. Dunsterville 114 (Tyrr in Herb. Ames). Octomeria deltoglossa Garay sp. nov. Epiphytica, caespitosa, usque ad 8 em. alta; radicibus filiformibus, flexuosis, glabris; caulibus secundariis erec- tis vel paulo arcuatis, vaginis 8, arcte adpressis omnino obtectis, 2-5 cm. longis; folio carnoso, oblongo-ligulato vel oblongo-lanceolato, apice acuto vel acutiusculo, basim versus sensim angustato, 2-4 em. longo, 1-3 mm. lato; floribus aggregatis, diaphanis; sepalis inter se simillimis, ovatis vel oblongo-ovatis, acutis, 8-5 mm. longis, 1-1.5 mm. latis; sepalis lateralibus paulo obliquis; petalis sepa- lis simillimis sed paulo minoribus, 8-4.5 mm. longis, 1-1.2 mm. latis; labello valde carnoso, deltoideo, apice acuto et pulvinato-incrassato, 1.5-1.8 mm. longo, 1—1.3 mm. lato; columna crassa, arcuata; ovario pedicellato ca. 2 mm. longo. This new species is closely related to the Brazilian Octo- meria rhodoglossa Schitr., but differs from it in having smaller flowers and a dissimilar lip. Venezugta: Rio Carrao. G. C. K. Dunsterville 289 (Tyre in Herb. Ames).—Territorio Amazonas, Cerro Sipapo (Paraque), 1400 m. alt. B. Maguire & L. Politi 27727 (Herb. N.Y. Bot. Gard.). [ 199 ] PLaTE XXXVII nities & Pollino Column & Lip Cress section of taaf Column be Lip OcToMERIA DELTOGLOsSA Garay Drawn by G. C. K. DunstervILLe Malaxis tipuloides (Lindl.) O. Ktze. var. exigua Garay var. nov. A typo floribus multo minoribus differt. Planta usque ad 30 cm. alta; foliis 5, basilaribus, aggregatis, ovato- lanceolatis, acutis vel subacuminatis, 12-13 cm. longis, 1-1.5 em. latis; racemo erecto, densissime multifloro; bracteis lanceolato-setaceis, pedicellis aequilongis; flori- bus parvulis; forma sepalorum et petalorum et labelli illa typi simillima; sepalis tantum 4-5 mm. longis. Whether to recognize this collection as a new species or merely as a variety of M. tipuloides was a matter of de- liberation for a long time. No morphological differences of specific value can be observed in the floral structures between this collection and the typical species. However, the small dimensions are so striking that I feel it neces- sary to establish it at least as a variety. Cotompia: Dept. Cundinamarca. Cordillera Oriental; Hato Grande, east side of Rio Muchindote, 13 km. east of Gachetd. 2600 meters alt. June 14, 1944, Martin L. Grant 9456. Collected under the aus- pices of the U.S. Foreign Economic Administration: Colombian Cin- chona Mission (Tyre in Herb. Ames; Dupticare type in U.S. Nat. Herb. ). Liparis serpens Garay sp. nov. Terrestris, 5-8 cm. alta; rhizomate satis gracili, flex- uoso, prorepenti, vaginis remotis, scariosis, cucullatis, acutis, 7-10 mm. longis obsesso; folio juxta basim pseu- dobulbi singulo, prominenter petiolato, lamina ovato- cordata, acuta vel subacuminata, basi abrupte rotundata, margine leviter crispata, petiolo incluso 8—5.5 cm. longa, 1.5-8 cm. lata; pseudobulbo vix evoluto, ca. 5-7 mm. longo; inflorescentia singula, erecta vel leviter arcuata, satis gracill, 4.5-8 cm. longa, dimidio superiore dense multiflora; bracteis ovato-lanceolatis, acutis, quam ovariis pedicellatis brevioribus, 83-4 mm. longis; sepalis ovatis, obtusis, margine revolutis, nervo mediano prominenti, [ 201 ] 4.5-5 mm. longis, 1.25 mm. latis; sepalis lateralibus leviter obliquis; petalis ligulatis, margine revolutis, uni- nerviis, 5 mm. longis, 0.8 mm. latis; labello a basi cune- ata subito dilatato, antice bilobo, in medio apiculato; lobis divergentibus oblongis, margine exteriore apicem versus serrulato-denticulata; disco in medio leviter in- LipaRis SERPENS Garay crassato, ad basim callo transverse semiquadrato, antice leviter excavato ornato, 6 mm. longo, in medio 8 mm. lato, inter lobos apicales 6 mm. lato; columna gracili, leviter arcuata, apice incrassata auriculataque, ca. 8 mm. longa; ovario cum pedicello alato, ca. 5 mm. longo. Liparis serpens is closely related to L. neuroglossa Rchb.f. and L. elegantula Kriinzl. It differs from the former in its trailing habit and in the shape of the lip. Liparis elegantula, which is very similar vegetatively, appears to be constantly bifoliate and to have a differ- ently shaped lip. [ 202 ] Cotomsia: Dept. Cundinamarca, Quebrada del Chicé, Bogota. Ter- restrial in shady woods. Flowers light green, lip bright green with violet veins. 2700-2800 meters alt. Martin Schneider 95 (Tyrr in Herb. Ames no. 68507.) Epidendrum superpositum Garay sp. nov. Epiphytica, ramosa, usque ad 40 em. alta; radicibus crassiusculis, glabris; caulibus superpositis, erectis vel paulo arcuatis, vaginis amplis omnino obtectis, apice 2-3-phyllis; foliis ovatis, acutis, satis coriaceis, 3-6 cm. longis, 0.8-1.4 em. latis; pedunculo abbreviato, 1-2 cm. longo; racemo densifloro, subeapitato, recurvo; bracteis setaceis ovariis brevioribus; sepalo postico obovato, apice acuto, 8-nervio, 8-10 mm. longo, 3-4 mm. lato; sepalis lateralibus obliquis, obovato-falcatis, apice cuspidatis, 3- nerviis, 8-10 mm. longis, 4-6 mm. latis; petalis lineari- obovatis, obtusis, 1-nerviis, 8-10 mm. longis, 1-1.5 mm. latis; labello trilobo, lobis lateralibus late rotundatis, margine leviter undulatis, lobo terminali antico, in medio exciso, sic bilobulato, disco basi bicalloso, 5-8 mm. longo, 10-12 mm. lato; columna labello adnato; ovario cum pedicello ca. 1.5 cm. longo. This new species is apparently similar in appearance to Epidendrum obliquum Schitr., 17. soratae Rehb.f. and E.. Steinbachii Ames, but it differs from them in various floral structures, especially in the shape of the lip. Venezue_a: Junquito. G. C. K. Dunsterville 220 (Tyrer in Herb. Ames no. 68942). Hexadesmia Dunstervillei Garay sp. nov. Epiphytica, caespitosa vel prolifera, usque ad 20 cm. alta; radicibus filiformibus, flexuosis, glabris; pseudo- bulbis longe stipitatis, cylindraceis, bifoliatis, usque ad 6 cm. longis; foliis ellipticis vel oblongo-ellipticis, utrin- que attenuatis, apice acutis, 2-7 cm. longis, 0.7—2 cm. latis; inflorescentia satis abbreviata, pauci- (1—-2-) flora: [ 203 ] PLATE XXXVITI Lateral Sepal 220 EprpeNDRUM sUPERPOSITUM Garay Drawn by G. C. K. Dunstervitir FUATE. XX XTX Gcxo 256 \ wi : 7 a i= Anther & Pollina 336 HexapesMiA Dunstrervit_et Garay Drawn by G. C.K. DunsrervILie bracteis ovato-lanceolatis, acuminatis, ca. 2.5 mm. longis; sepalis lanceolato-ovatis, acuminatis, 5—7-nerviis; sepalo postico 15-16 mm. longo, 4-6 mm. lato; sepalis lateral- ibus leviter obliquis, 15-16 mm. longis, 4-5 mm. latis: petalis ovato-lanceolatis, acuminatis, 15-16 mm. longis, 6-7 mm. latis; labello subquadrato-fiabellato, apice acuto, disco bicarinato, 14-15 mm. longo, 8-9 mm. lato; columna leviter arcuata, basim versus paululo dilatata et in pedem brevissimum producta. Hewxadesmia Dunstervillei closely resembles H. Jime- nesu Schltr. from Costa Rica but is easily separable from that taxon by the shape of the lip. This new species is dedicated to my good friend G. C. Kk. Dunsterville, Pres- ident of the Shell Oil Co. in Venezuela, with whom I am preparing an iconography of the Orchids of Venezuela. VenezurLa: Estado Miranda, Santa Teresa, Guatopo. Dec. 31, 1955. G.C. K. Dunsterville 336 (Tyrr in Herb. Ames). Bifrenaria minuta Garay sp. nov. Kpiphytica, caespitosa, minuta, usque ad 12 em. alta; radicibus filiformibus leviter flexuosis, glabris; pseudo- bulbis aggregatis, ovoideis, unifoliatis, 1.5—2 cm. altis; folio pergameneo, subplicato, apice acuto, basi in petio- lum 2-8 cm. longum attenuato, petiolo incluso 10-12 em. longo, 1-1.5 em. lato; inflorescentia juxta basim pseudobulborum nata, uniflora; pedunculo capillari, ca. 1-1.5 cm. longo; sepalo postico ovato-oblongo, apice acuto, 5-nervio, 9 mm. longo, 4.5 mm. lato; sepalis lat- eralibus oblique ovatis vel ovato-oblongis, apice acutis, 5-nerviis, 10 mm. longis, 6 mm. latis; petalis trulliformi- bus apice acutis, 8-nerviis, 8.5 mm. longis, 8 mm. latis; labello rhombeo, antice truncato, disco vix calloso, 10 mm. longo, 6 mm. lato; columna leviter clavata, basi in pedem producta; ovario cum pedicello ca. 6 mm. longo. This new species is quite unique in the genus on ac- [ 206 ] PLATE XL Anther Column & Lip. BiIrRENARIA MINUTA Garay Drawn by G. C. K. DunsTervILLe count of the extremely small flowers. It is apparently a borderline species between Bifrenaria and Mawillaria. Venezur.a: Rio Carrao. G. C. K. Dunsterville 262 (Tyrer in Herb. Ames. ) Bifrenaria grandis (A7riénzl.) Garay comb. nov. Lacaena grandis Kriinzl. in Fedde Repert. 25 (1927) 25. An examination of an isotype (Buchtien 501) in the U.S. National Herbarium reveals that Kriinzlin’s con- cept isa member of the genus Bifrenaria, section Lind- leyella, and is closely allied to the Colombian B. picta (Schltr.) Schweinf. Bifrenaria grandis is native of Bolivia. Maxillaria equitans (Sch/ltr.) Garay comb. nov. Camaridium equitans Schltr. in Fedde Repert. Beih. 7 (1920) 176. Marsupiaria equitans Hoehne in Arquiv. Bot. Est. S. Paulo 2, pt. 4 (1947) 71. A recent collection (Cuatrecasas 22092) from the De- partamento del Valle in Colombia shows that this spe- cies has somewhat falcate leaves, similar to those of MM. vandiformis Schweinf., a character not mentioned by Schlechter in the original description. Since Camaridium is not tenable, the above transfer is proposed. Maxillaria Sophronitis (Rchb.f) Garay comb. nov. Ornithidium Sophronitis Rchb.f. in Bonpl. 2 (1854) 18. This rather showy species has not previously been transferred to the genus Maaillaria. It is known from Venezuela only and has recently been collected by Mr. Dunsterville (Dunsterville 388). Scelochilus corydaloides (A7iéinzl.) Garay comb. nov. [ 208 | Rodriguezia corydaloides Kriinzl. in Fedde Repert. 25 (1928) 24. An examination of the type (Buchtien 528) indicates that this taxon is referable to the genus Scelochilus. Rodriguezia Teuscheri Garay sp. nov. Epiphytica, caespitosa, ca. 12 em. alta; radicibus flex- uosis, glabris; pseudobulbis ovoideis, lateraliter com- pressis, unifoliatis, 2-8 cm. longis; folio coriaceo, ellip- tico vel oblongo-elliptico, canaliculato, apice acutiusculo, basi in petiolum brevem producto, 8-10 cm. longo, 2.5-8 em. lato; inflorescentia juxta basim pseudobulborum nata, erecta vel paulo arcuata, pauciflora, ca. 8 cm. longa; bracteis ovatis, acutis, ca. 5 mm. longis; sepalo postico valde concavo, obovato, apice obtusiusculo, 1.5 cm. longo, ca. 1 em. lato; sepalis lateralibus usque ad apicem connatis, reflexis, coneavis, falcatis, ca. 2.5 em. longis; petalis obovato-spathulatis, apice obtusis vel subtrunca- tis, 1.5 em. longis, 1 cm. latis; labello in ambitu trape- zoideo, antice emarginato, in medio sinuato, basi in pro- cessum carnosum producto, disco per totam longitudinem lamellis 2, parallelis ornato; lamina labelli ca. 2 cm. longa, antice 1.5 cm. lata; columna clavata, satis gracili, ca. 1.7 cm. longa; ovario cum pedicello ca. 2.5 cm. longo. The specific name is given in honor of Dr. Henry Teuscher, Curator of the Montreal Botanical Garden, Canada, whose keen interest inand knowledge of orchid- ology is widely known and recognized. Ecuapor: 100 km. from Guayaquil. H. Teuscher s.n. (Type in Herb. Ames). Flowered in cultivation at the Montreal Botanical Garden. Cryptocentrum Lehmannii (2chb.f.) Garay comb. nov. Aéranthus (Cryptoplectri) Lehmanni Rehb.f. in Otia Bot. Hamb. (1878) 10. [ 209 ] EXPLANATION OF THE ILLUSTRATION Pirate XLI. Habit photograph of Ropricurzia Teuscurert L, 4, Garay. This is a photograph of the type material as cultivated in the Montreal Botanical Garden, Canada. Photograph by Romro MrLocur [210 | PLATE XLI Campylocentrum Lehmanni (Rehb.f.) Schltr. in Fedde Repert. Beih. 8 (1921) 164. An excellent record of the type of the Hcuadorian Aéranthus Lehmanni in the Ames Herbarium reveals that this concept is undoubtedly referable to the genus Cryptocentrum. Apparently C. Jamesonu Benth, is the only other species of the genus which has been reported from Keuador. Although these two taxa seem to be very similar vegetatively, C. Lehmanni may be separated by the presence of six to seven sheaths on the peduncle in contrast to one or two in C. Jamesonu, as well as by the pandurate and long-acuminate lip in the former, whereas it is ovate in the latter. In his system of the Orchidaceae, Schlechter assigned the genus Cryptocentrum to the subtribe Maaillarieae. In view of the peculiar vegetative and floral structures, however, I cannot agree with this opinion. We find here a type of vegetative growth closely resembling a mono- podial development. The leaves are arranged distichously on a shortened stem, but only a few terminal ones seem to be persistent, at least during anthesis, while the lower part of the stem bears the remnants of the sheathing bases of the leaves of previous years. ‘The one-flowered peduncles are always borne in the axils of these sheath- ing bases on the lower portion of the stem. This condi- tion is observable in every species assigned to Cryptocen- trum, with the exception of C. pseudobulbosum Schweint. In C. pseudobulbosum, we find, in addition, a well devel- oped pseudobulb, quite an unusual condition, but, from the systematic point of view, exceedingly important. With the evidence from C. pseudobulbosum, it is obvious that this genus has to be kept among the genera with sympodial growth. It may be mentioned as a mere curi- osity that this vegetative growth is entirely homologous with that of the old world genus Phreatia. C212) In the floral structure of Cryptocentrum, furthermore, we find evidence of a more complex nature than would be normal for the subtribe Maaillarieae. The lateral se- pals are connivent at the base and form a very prominent sepaline spur to which the claw of the lip is internally adnate for its entire length. This feature is observable in the genus Sepalosiphon in addition to Cryptocentrum. (Sepalosiphon is known only from New Guinea.) Though we have here a striking homology in floral structures, the two genera are vegetatively very dissimilar. In both cases the column is very short, more or less petaloid and with- out a foot. The incumbent anther rests on a more or less ascending and acicular rostellum. In applying these observations to Schlechter’s ‘‘Sys- tema Orchidacearum,’’ we find that the characters of the subtribe Mawillarieae are very different, as shown in the following tabular comparison. MAXILLARIFAE CRYPTOCENTRUM Rostellum not evident. Rostellum prominent. Column elongate and with a definite Column short and footless. foot which forms, with the lateral se- No mentum present. pals, a more or less prominent mentum. Lateral sepals not spurred. Lateral sepals spurred. Lip articulate with the column foot; Lip adnate to the spur; disc dise with a longitudinal callus. ecallose. It is clear that Cryptocentrum cannot be maintained in the subtribe Maaillarieae. In following through Schlech- ter’s system, the characters of Cryptocentrum as tabu- lated above, give us substantial ground to warrant the erection of a new subtribe, which should be placed be- tween Ornithocephaleae and Saundersieae. Cryptocentreae Garay subtribus nova. Pollinia 4. Plantae epiphyticae, pusillae. Pseudobulbi rudimentarii vel deficientes. Caules satis abbreviati, pseu- [ 213 | domonopodium formantes. Folia disticha. Inflorescen- tiae uniflorae. Sepala calcarata. Labellum sepalorum cal- cari intus adnatum. Columna apoda. Rostellum promi- nens, aciculare. Stipes polliniorum abbreviata. Genera: Cryptocentrum Bentham. Anthosiphon Schlechter. Notylia Norae Garay sp. nov. Epiphytica, pusilla, ca. 2.5 em. alta; radicibus filiform- ibus, flexuosis, glabris; foliis equitantibus, oblique ovato- lanceolatis, carnosis, 5-8 mm. longis, 1.5-2 mm. latis; inflorescentia erecta vel paulo arcuata, pauciflora; brac- tels ovatis, acutis, ca. 1 mm. longis; floribus diaphanis, lilacinis ; sepalo postico ovato, conduplicato, apice acuto, 1-nervio, 3 mm. longo, 1.7 mm. lato; sepalis lateralibus liberis, anguste oblongo-lanceolatis, acutis, 1-nerviis, 4 mm. longis, 0.8 mm. latis; petalis ovato-lanceolatis, acutis, 1-nerviis, 4mm. longis, 1 mm. latis; labello longe unguiculato, deinde in laminam ovatam in medio leviter constrictam expanso, apice acuto, basi minutissime cili- ato, 3-nervio, 8.5 mm. longo, antice 1.8 mm. lato; col- umna erecta, capitata, ca. 2 mm. longa; ovario cum pedicello ca. 2.5 mm. longo. Notylia Norae is related to the Bolivian N. hlacina Kriinzl. but differs from it in the shape of the petals and lip. It gives me great pleasure to dedicate this beautiful miniature orchid to Mrs. Dunsterville, whose inspiration and special interest in botanicals has been of indispensa- ble help in the preparation of the Lconography of the Venezuelan Orchids. Venezurta: Kavanupa, Rio Uaiparu. On small tree near water’s edge. G. C. K. Dunsterville 442 (Tyre in Herb. Ames). Telipogon Teuscheri Garay sp. nov. ($ Brevicaules). Radicibus satis longis, copiosis ; cauli- [ 214 | PiateE XLII 2mm Column”, Anther & Potliua Cotumn 4 Up 442 Noryiia Norar Garay Drawn by G. C. K. DunsterviLie EXPLANATION OF THE ILLUSTRATION Pirate XLIII. Terreocon Teuscueri Garay. 1, plant, two thirds natural size. 2, flower, expanded, slight- ly enlarged. 8, lip, twice natural size. Drawn by Evmer W. Situ [ 216 | PiateE XLIII TELIPOGON Teuscheri yaray bus abbreviatis, ca. 4-foliatis; foliis satis carnosis, oblongo- lanceolatis, acutis, basi articulatis, 2.5—4 cm. longis, 4-8 mm. latis; inflorescentiis erectis, vel leviter flexuosis, quam foliis duplo longioribus, apice 2—4-floris; pedunculo compresso, alato, usque ad 7 cm. longo; bracteis ovato- lanceolatis, dorso leviter carinatis, ovariis duplo breviori- bus; floribus in genere inter mediocres; sepalis ovato- triangularibus, acutis, 3-nerviis, dorso prominenter car- inatis, 14 mm. longis, supra basim 6 mm. latis; petalis rhombeo-ovatis, acutis vel subacuminatis, 11-nerviis, 16 mm. longis, 13 mm. latis; labello transverse subrotundo, acuto, in medio breviter apiculato, 17-nervio, 138 mm. longo, 17 mm. lato; disco basim callo deltoideo, hirsuto ornato; columna humili, hispido-setacea; ovario gracili- ter pedicellato, cum pedicello ca. 2 cm. longo. Telipogon Teuscheri differs from T. semipictus Rchb.f. in having a well developed callus at the base of the lip and somewhat smaller flowers. Velipogon Wallisii Rchb.f. is very similar to this new species, but its flowers are smaller and its petals are only 7-nerved. Ecuapor: Santa Rosa. 11,000 ft. alt. June 1956. H. Teuscher 12 (Tyrer in Herb. Ames). [ 218 ] TWO ADDITIONS TO THE ORCHID FLORA OF VENEZUELA BY CHARLES SCHWEINFURTH Dunrinec the course of my investigations on the orchid flora of the Guiana Highlands with special emphasis on Venezuela, the following two species which seem to be amply distinct were discovered. Vargasiella venezuelana C. Schweinfurth sp. nov. Herba terrestris, elata, robusta, in parte inferiore de- cumbens. Caulis vaginis numerosis, adpressis, imbrican- tibus, foliiferis omnino celatus. Folia elliptica vel elliptico- oblonga, valde acuminata, ad basim articulatam longe attenuata, submembranacea, plicata. Inflorescentia e folii medii axilla exoriens, elata; pedunculus vaginis nonnullis brevibus, tubulatis remote ornatus; racemus perlaxe pauciflorus. Flores mediocres, laete purpurei, subcrassi, cum sepalis longitudinaliter concavis. Sepalum dorsale expansum lanceolato-oblongum, obtusum vel acutum. Sepala lateralia valde similia sed paullo majora, obliqua. Petala sepalis multo breviora et latiora, ovalia vel oblongo-elliptica, obtusa vel subacuta. Labellum simplex, unguiculatum; unguis brevis sed distinctus, a basi oblonga abrupte dilatatus, carina mediana crassa, apice divisa ornatus; lamina leviter recurva, triangulari- ovata, apice rotundata, basi prominenter cordata. Col- umna crassa, brevis. [ 219 ] EXPLANATION OF THE ILLUSTRATION Prate XLIV. VarGAsIELLa VENEZURLANA C, Schweinf. 1, plant, one fourth natural size. 2, flower from side, natural position, one and one third times natural size. 3, lip and column from side, natural position, twice natural size. 4, petal, twice natural size. 5, lip, natural position, twice natural size. Drawn by Etmer W. Situ [ 220 ] PuateE XLIV a VARGA SIELLA_ Ri C. Schweinfurth_ 2Y venezuclana Plant terrestrial, tall and robust, with a decumbent, sparsely rooting stem, up to about 187.5 cm. long in- cluding the erect inflorescence. Roots very remote, soli- tary, fibrous, rather stout, finely pubescent. Stem en- tirely concealed by appressed, imbricating, evanescent, leaf-bearing, tubular sheaths of which the lower ones are scarious and disintegrating into fibres and the middle and upper ones are green and leaf-bearing. Leaves elliptic or elliptic-oblong, acuminate to a sharp point, long- narrowed below to an articulated base, submembrana- ceous, plicate, with five to seven prominent nerves, about 14-17 cm. long, up to 8.8 cm. wide. Inflorescence spring- ing from the axil of one of the middle leaves, up to 52 em. high; peduncle dull lavender, glabrous, about 48 cm. or less long, remotely provided with about six short, tubular, acute sheaths; raceme very loosely few- (about 9-) flowered, the rachis about 9 cm. or less long. Floral bracts small, narrowly lanceolate, concave, equaling about half of the pedicellate ovary which is up to 2.3 cm. long. Flowers rich purple, subfleshy, with the sepals projecting backward and the petals erect. Sepals longitu- dinally concave. Dorsal sepal when expanded lanceolate- oblong, obtuse or acute, about 1.5 cm. long and 4.4 mm. wide. Lateral sepals similar, obliquely lanceolate-oblong, obtuse or acute, about 1.6 cm. long and 6 mm. wide. Petals distinctly shorter and broader than the sepals, oval to oblong-elliptic, obtuse to subacute, somewhat concave, up to 1.2 cm. long and 7 mm. wide, the margins often somewhat irregular. Lip simple, clawed, with involute, undulate margins; claw short but distinct, abruptly di- lated from an oblong base, about 3 mm. long, provided with acentral fleshy keel that divides abruptly into two diverging branches; lamina gently recurved, triangular- ovate, rounded at the apex when expanded, conspicu- ously cordate at the base, about 1.1 cm. long in the cen- [ 222 | ter and up to 1 cm. wide; dise fleshy-thickened in the middle below. Column stout, about 5 mm. high. This plant is the second species referable to the remark- able South American genus Vargasiella. It is clearly dis- tinct from V’. peruviana C. Schweinf. in its greater vege- tative proportions, its bright purple (rather than whitish pink) flowers, its narrower, longitudinally concave sepals and its distinctly clawed lip. VenezueL_a: State of Bolivar, Chimantd Massif, northwestern part of summit of Abadcapa-tepui, at 2125-2300 meters altitude, in Bonnetia forest, April 13, 1953, Julian A. Steyermark 74914 (Tyre in Herb. Ames No. 69272). Oncidium vagans C. Schweinf: sp. nov. Herba magna, vagans, terrestris vel scandens, valde variabilis. Rhizoma cauliforme, plusminusve crassum, simplex vel in nodo ramosum, vaginis arctis, tubularibus, scarlosis, evanidis omnino celatum, radices remotas, sim- plices, longas nune emittens. Pseudobulbi remoti, cylin- dracei vel anguste pyriformes, prope apicem saepissime bifoliati, vaginis nonnullis, scariosis, evanidis inferne or- nati. Folia lanceolata vel anguste oblongo-lanceolata vel anguste elliptico-lanceolata, acuta et apiculata, papyra- cea. Inflorescentia folia multo superans, e pseudobulbi basi exoriens, superne perlaxe paniculata, suberecta; pe- dunculus vaginis nonnullis, parvis, arctis ornatus; rhachis plusminusve fractiflexa, cum ramis horizontalibus vel de- curvis laxe paucifloris. Flores parvi, cum segmentis late patentibus. Sepalum dorsale perbreviter unguiculatum, oblongo-ovatum vel oblongo-ovale, acutum vel apice ro- tundatum, trinervium. Sepala lateralia oblique obovato- oblonga vel oblongo-oblanceolata, apice rotundata vel acuta, quadrinervia, ad basim paululo unguiculatam an- gustata. Petala breviter unguiculata, abrupte ovato- oblonga vel oblongo-ovalia, apice lato plusminusve retusa et saepe subacuta, quinquenervia. Labellum segmentis [ 223 ] EXPLANATION OF THE ILLUSTRATION Pirate XLV. Onciprum vaaans C. Schweinf. 1, plant, one fourth natural size. 2, flower, natural position (lip expanded), twice natural size. 3, dorsal sepal, two and one half times natural size. 4, petal, about twice natural size. 5, lateral sepal, twice natural size. 6, column, from side, three times natural size. 7, leaf (of another collection), approximately one half natural size. Drawn by Rutu Barton [ 224 J PLatTE XLV ONCIDIUM VACaANS x Schweinf ceteris multo majus, panduratum vel trilobatum; lobi laterales semiovato-triangulares cum angulis exterioribus late rotundatis; lobus anterior reniformi-flabelliformis, apice latissime rotundato emarginatus; discus callo bi- seriato, basi cuneato, crasse pluridentato ornatus, cum parte apicali conspicue tridentata. Columna parva, cum alis latis, subquadratis superne ornata et crassificatione divaricatim bilobata inferne praedita. Plant large, sprawling, very variable, terrestrial or climbing. Rhizome stem-like, more or less stout, simple or virgately branched at one of the nodes, entirely con- cealed by close, tubular, scarious, evanescent sheaths, occasionally producing at remote intervals long, simple, fibrous roots. Pseudobulbs about 3-13 cm. apart, cylin- dric to narrowly pyriform, commonly 2- (rarely 3-) leaved near the summit, about 2-6.4 cm. long, provided below with several narrow, scarious, evanescent sheaths of which one is produced into an oblong-lanceolate blade. Leaves lanceolate to narrowly oblong-lanceolate or narrowly elliptic-lanceolate, acute and apiculate, papyraceous, 4— 14.6 cm. long, up to 1.6 em. wide. Inflorescences much surpassing the leaves, arising from the base of the pseudo- bulb, very loosely paniculate above, about 44.5-117.5 cm. long, suberect, often diffuse above; peduncle pro- vided with several remote, small, inconspicuous sheaths, up to 73.5 cm. long; rachis more or less fractiflex with the horizontal or decurved branches loosely 8- to 7- flowered above. Flowers small, with widespreading seg- ments, pale greenish or cream-colored to pale yellow with irregular brownish or purple bands. Dorsal sepal shortly and narrowly clawed, oblong-ovate or oblong-oval, acute to rounded at the apex, 8-nerved, about 6—6.8 mm. long and 3.2—4.5 mm. wide. Lateral sepals obliquely obovate- oblong or oblong-oblanceolate, rounded to acute at the apex, 4-nerved, narrowed to an indistinctly clawed base, [ 226 ] about 8-11 mm. long and 3-4 mm. wide. Petals from a short claw rather abruptly ovate-oblong or oblong-oval, more or less retuse and often subacute at the broad apex, often slightly narrowed or indented on each side, about 6.38-9 mm. long and 4-6 mm. wide, indistinctly 5-nerved. Lip much larger than the other segments, pandurate or 3-lobed, white to cream-colored with purple spots and a yellow callus, about 11-13 mm. in greatest length, about 8-8.7 mm. wide across the basal portion; lateral (basal) lobes semiovate-triangular, broadly rounded at the outer angles, with a more or less distinct, semiorbicular lobule on each side of the base; anterior lobe reniform-flabellate, emarginate at the broadly rounded apex, about 10-11.5 mm. wide; disc with a prominent callus below consist- ing of two series of which the lower one is cuneate at the base and composed of several small, fleshy teeth and the anterior one of three larger, fleshy teeth; on either side of the callus there is sometimes (as in the type) a single fleshy tooth. Column small, about 3.5 mm. high, with a pair of broad, porrect, subquadrate wings above and a fleshy, divaricately bilobed thickening below. Oncidium vagans appears to be allied to the Brazilian O. Warmingu Reichb.f., but differs in having entire, differently shaped lateral lobes of the lip, sessile mid-lobe and subquadrate column-wings. VENEZUELA: Bolivar, Central Section, Chimanta Massif, 2120 meters alt., swampy depression in wet savanna along east branch of head- waters of Rio Tirica, February 12, 1955, ‘‘Ivs coriaceous, olive; pseu- dobulbs subterete. Peduncle reddish brown; lip white to cream with purple spots in center and at apex; callosity yellow with purplish spots ; lateral lip appendages white; column purplish above, within whitish with greenish yellow at base; sepals and two lateral petals pale yellow to cream with broad irregular purple bands, locally frequent around hammocks,’’ Julian A. Steyermark & John J. Wurdack 762 (Tyrer in Herb. Ames 68534.—Amazonas, Cerro Duida, Rio Cunucunuma (Culebra Creek drainage), 1600 meters alt., ‘‘terrestrial; outer peri- anth members pale greenish, pale brown mottled ; lip white with mot- ea tled appendages and yellow protuberance. In interzone between Bon- netia and cloud forests,’? November 19, 1950, Bassett Maguire, R.S. Cowan & John J. Wurdack 29553.—Bolivar (The Phelps Uaipan-Tepui Expedition), at the summit, 1700 meters alt., flowers white, January 25, 1948, Kathleen D. Phelps & Charles B. Hitchcock 358.— Bolivar, 1800 meters alt., ““semitrepadora o erguida en formacién arbustiva baja. Parte central del plat6é de Auyantepui, lugar soleado y seco,’’ April 1956, Vareschi & Foldats 4933.—Same date as the last, Foldats 2599,—Cerro Auyantepui, at the summit, Alto Caroni, 2100 meters alt., January 1949, F. Cardona 2743, 2744. [ 228 ] A LITTLE-KNOWN CULTIVATED PLANT FROM NORTHERN SOUTH AMERICA BY RicHarp Evans SCHULTES I'r is not often that an important economic plant, culti- vated over a wide area, hides out from the eyes of plant- explorers, anthropologists, missionaries and travellers— even in an area with such a poorly understood agricul- ture as the Amazon Valley. Yet that has apparently come to pass in the case of Solanum Topiro HBA. I During my travels in the northwestern part of the Amazon basin, principally in Colombian territory, it was my custom always to study the plants cultivated by the Indian inhabitants of the region. It is possible, in general, to arrange the plants which the natives of this tropical forest-region grow into two large categories on the basis of the manner in which they are cultivated. Native agriculture in the western Ama- zon has been called, and [ think perhaps erroneously, ‘*primitive.’” Whether or not it be primitive, it is true that no machinery is employed, and little or no formal planning underlies agricultural practices. Plants are either set out in fields devoted to a single crop or else they are put in or allowed to spring up individually at random mingled with several or many different species. The first of these two categories can claim, at least, in the northwestern part of the vast Amazon, only two spe- cies: yuca (Manihot esculenta Crantz) and coca (diry- throvylon Coca Vam.). All other cultigens grow singly [ 229 ] or in groups of several individuals scattered through fields of yuca or coca, along the edges of these fields, in clear- ings or close to house-sites. Many species of economic plants fall into this second category. We might cite, as a few examples, the following: Phyllanthus spp., Teph- rosia tovicaria Pers. and Clibadium asperum (Aubl.) DC. (fish-poisons); Nicotiana Tabacum LL. and Banisteriopsis spp. (narcotics): Capsicum frutescens LL. (spice): and Herrania nitida (Poepp.) R. E. Schult. (food). In wellnigh every Indian agricultural plot in the Co- lombian Amazonia, especially along their margins and in the immediate vicinity of dwellings, two shrubby spe- cies of Solanum are grown for their edible fruits. The more abundant of these has been determined as Solanum Topiro. We have still not been able satisfactorily to iden- tify the second, and, common though it be in the areas of Indian agriculture, it may well represent a concept hitherto undescribed. The present paper, however, treats only of Solanum Topiro. It is with pleasure that I acknowledge the very help- ful interest of Mr. and Mrs. Kendal Morton of the Mor- ton Collectanea (University of Miami) in my attempts to identify specimens of this plant which I had brought back from Colombia. I have also here to thank Mr. Klmer W. Smith, artist at the Botanical Museum of Harvard University, who has seen So/ano Topiro in its native state in garden-plots in the Colombian Vaupés, for his outstanding drawing which was made possible through a grant from the National Science Foundation. This is the first time that So/anum Topiro has been illustrated. I] It may be well to outline what little we know of the history and recent introduction into horticulture of this species, [ 230 | A scrutiny of the major works on tropical fruit plants indicates that Solanum Topiro has been completely neg- lected and probably unknown as acultigen. It does not, for example, appear in Wilson Popenoe’s ‘‘Manual of tropical and subtropical fruits’* (1920), in F. C. Hoehne’s ‘*Frutas indigenas’’ (1946) nor in Adolfo Ducke’s **Plan- tas da cultura precolombiana na Amazonia brasileira. . .” in Bol. Téen. Instit. Agron. Norte, no. 8 (1946). Fur- thermore, standard anthropological and botanical papers on Indian agriculture in the American tropies fail to mention it: Robert H. Lowie, *‘The tropical forests: an introduction” and Carl O. Sauer, ‘‘Cultivated plants of South and Central America” in Handbook of South American Indians [ed. J. H. Steward], Bur. Am. Ethnol. Bull. 143, 8 (1948) 2 and 6 (1950) 487, respectively. As Fennell (Fennell, Joseph L. ‘*Cocona—a desirable new fruit’’ in For. Agric. 12 (1948) 181) has written: ‘*To what extent, if any, the cocona [Solano Topiro] may have reached the gardens of the outside world is difficult to say. That it appears even now to be essentially un- known to horticulture leads me to believe, in light of its impressive appearance and apparent usefulness, that it may never have previously lett its secluded habitat as a recognized fruit of value.” So faras I have been able to ascertain, the first serious attention paid by botanists to Solano Topiro as a culti- gen dates from the middle of the 1940’s. During this period, seeds of the plant were collected **from the little- explored reaches of the upper Amazon”’ (presumably in Peru) and established in the Experiment Station at Tingo Mariain Peru. Eventually, it was introduced to the In- stituto Interamericano de Agricultura Tropical in Tur- rialba, Costa Rica (Fennell, loc. cit. ; Ochse, J. J.‘ Sola- num hyporhodium or cocona’” in Proc. Fla. State Hort. Soc. 66 (1958) 211) from which centre it began to attract horticultural attention. [ 281 | The vernacular name for Solanum Topiro in Peru is reported to be cocona. This is borne out by notes on her- barium specimens (Aillip & Smith 273607, 27323) collected inthe Amazonian part of Peru in 1929. This convenient epithet followed the plant in its several introductions and has now, in the literature, been accepted as a standard common name. Unfortunately, however, cocona has been erroneously identified and has, in agricultural institutions as well as in the scientific and popular literature (Iennell, loc. cit.; “‘Cocona™” (abstract of foregoing article) in Keon. Bot. 8 (1949) 216; Ochse, loc. cit. : [ Bischoff, William] ‘'Phe Peach Tomato” in Miami Daily News (March 7, 1954) ) been determined as Solanum hypor- hodium A. Br. et Bouche. It has not been possible for me to ascertain where this erroneous identification was made, nor do | find in any of our larger herbaria a specimen-voucher upon which it could have been based. In the card-file at the Subtropical kxperiment Station in Homestead, Florida, the follow- ing data relative to cocona are to be found: **Solanwm hyporhodium A. Br. & Bouché. Native to Australia. In 1948, seeds were received from F. B. Harrington, Natal: all plants dead by 1950 from nematodes. ° That there must have been a sudden and widespread distribution of seeds of this species is evidenced by the fact that the Subtropical Experiment Station, believing the plant to be native to the Old World, made an in- troduction trom South Africa. Ochse (loc. cit.) brought the species into cultivation at the University of Miami apparently directly from Costa Rica. In July, 1958, 1 found it growing experimentally at the Imperial College of ‘Tropical Agriculture in Trinidad, the result of a rel- atively recent introduction but with no record of the provenience of the seed. Identification of the material of Solanum Topiro which [ 282 ] I had gathered in the northwest Amazon puzzled me for a number of years. I was finally able to match my ma- terial collected in Colombia under the name /u/o with cocona through illustrations of cocona found in the Mor- ton Collectanea and to learn that cocona had been deter- mined as Solanum hyporhodium. In trying to check the accuracy of this determination, I discovered that there was no material representative of Solanum hyporhodium in our American herbaria. The type, and, apparently the only specimen of this species, was preserved in the Berlin Herbarium and, of course, is no longer extant. There is, nevertheless, a photograph of the type of Solanum hy- porhodium in the Gray Herbarium. It is at once obvious that cocona cannot be referred to Solanum hyporhodium, for the photograph indicates that there are differences in the leaf and that iS. hyporhodium has stems and petioles which are heavily spinose. The cocona collected originally in Peru, as well as all of my own specimens from Colom- bia and recent Venezuelan material, is entirely without spines and has the petioles clad with a soft mottled indu- mentum. Fennell’s published notes (loc. cit.), indeed, stress the lack of spines. It was possible for me finally to identify cocona or lulo by using one of the common Venezuelan names of the species. The non-Indian population of the Colombian Amazonia know the plant as /w/o, because of its strong resemblance to the highland Solanwmn quitoense Lam., which, in the Andes of Colombia, is called /u/o. Along the border between Colombia and Venezuela, however, the non-Indian inhabitants refer to the plant by its Ven- ezuelan name fopiro or fupirvi. In H. Pittier’s **Manual de las plantas usuales de Venezuela” (1926) 885, there is an entry under the common name topiro: **Solanum Topiro Dunal, Synops. 10. 1810. Sin. fupiro. Especie herbacea, tomentosa, inerme, las hojas ovales, mis o [ yo mm t DoD | menos angulosas, las flores verduscas en glomérulos opuestos a las ultimas y los frutos ovoideos, grandes y comestibles. Es del Alto Orinoco.” Sturtevant (*‘Stur- tevant’s notes on edible plants’’ [ed. U. P. Hedrick}, (1919) 545) had reported Solanum Topiro,a species grow- ing on the ‘‘banks of the Orinoco,” as having an edible fruit known as the ‘‘turkey berry,” attributing his sources to G. Don Hist. Dichl. Pls. 4 (1888) 410. Anexcellent photograph of the type of Solanum To- piro trom the Paris Herbarium enabled me to authenti- cate the identification. The type, from the Humboldt and Bonpland Herbarium, was collected at San Fernando de Atabapo on the Rio Orinoco, Venezuela (AZumboldt et Bonpland 918). Vhere is, further, in Paris a reproduc- tion of an unpublished drawing of Solanum Topiro (No. 192) for Dunal’s Solanaceae, ed. 2. There can be no doubt that cocona from Peru and the /i/o from Amazon- ian Colombia refer to the same species as fopiro in Vene- zuela and that all three are So/anuwm Topiro. We are now able to offer an amplified description and extended dis- tribution for this species, as well as sundry ethnobotan- ical notes appertaining to its cultivation and use. Solanum Topiro Humboldt, Bonpland & Kunth ev Dunal Sol. gen. aff. syn. (1816) 10. ORIGINAL DESCRIPTION : 37. S. Topiro. S. caule herbaceo tomentoso, foliis ovato-oblongis sinuatis basi inaequalibus subtus leviter tomentosis supra sericeis, floribus aggregatis extra-axillaribus, baccis ovatis.— Humb, et Bonpl. ined.— Dun. Sol, Ed. 2 ined. tab, 921 f. 1 Hab. ad Orenocum (v.s.h. H. et B.). Shrub unarmed, robust, rank,up to about 5 feet (1.5m. ) tall. Branches stout, terete, scurfy-pubescent, grey-green in life. "Twigs densely white-stellate-tomentose. Leaves coarsely membranaceous, ovate in outline, at maturity [ a4 | up to about 48 cm. long, 86 cm. wide, basally inequi- laterally truncate, apically abruptly acute, marginally very distantly and very deeply sinuate, strongly petio- late (petiole up to 12 cm. long, densely white-stellate- tomentose); upper surface coarsely tomentose and densely clad with somewhat stiff sericeous white hairs (some of which have basally a stellate formation) sparsely interspersed with white stellate hairs: nether surface very softly and densely white-stellate-tomentulose; veins strong and conspicuous above and below, extremely densely white-stellate-tomentose. Inflorescence a lateral, very short-peduncled, few-flowered cyme. Flowers pedi- cellate; pedicels up to 6 mm. long, 1.5—2 mm. in diam- eter, very densely stellate-tomentulose. Calyx lobes somewhat crassulent, more or less triangular-ovate, api- ‘ally subacute, up to about 15 mm. long, very densely and softly greyish brown-stellate-tomentose without, glabrous but with minute white scabs within. Corolla membranaceous, white or greenish white, lobes oblong- ovate, up to about 20 mm. long, apically subacute, gla- brous within, densely stellate-tomentulose without. An- thers yellow, erect, linear, about 9-10 mm. long. Style terete, 7-8 mm. long. Ovary globose, very densely long- white-sericeous. Fruit subglobose to ovoid, ripening orange-red, sometimes tomato-red, densely and minutely stellate-tomentulose (hairs easily caducous upon hand- ling), becoming subglabrous upon ripening. Pulp acid- ulous. Seeds very numerous, flat, oval in outline, 3-4 mm. long, 2.2.5 mm. wide, pale yellowish. CoLtompia: Comisaria del Amazonas, Rio Caqueta, vicinity of La Pedrera. “‘Fruit dark red-rust, hairy. Flowers white. Bush.*’ April 1944, R. FE. Schultes 5881.—Comisaria del Amazonas, Rio Loretoyacu. Altitude about 100 m. September 1946, R. F. Schultes §& G. A. Black S394,—Comisaria del Amazonas, Rio Apaporis, Soratama, near mouth of Rio Kananari. ““Cultivated. Fruit edible. Flowers whitish with yel- low anthers. Fruit ovoid, orange, covered with hair in unripe stage. Lulo. Tupiri.’* March 1951, R. E. Sehultes 12081.—Comisaria del EXPLANATION OF THE ILLUSTRATION Prare XLVI. Soranum Topiro HBK, 1, flowering branch, with medium-sized leaf, about one half natural size. 2, portion of the upper surface of the leaf, greatly enlarged. 38, portion of the nether surface of the leaf, greatly enlarged. 4, inflores- cence, about one half natural size. 5, fruits, about one half natural size. 6, young leaf, about one half natural size. [ 236 ] PLATE XLVI SOLANUM. EXPLANATION OF THE ILLUSTRATION Prare XLVI. Fruit of Soranum Topiro represented by the collection Schultes 12081 from the Colombian Amazonas. Photograph by Ricuarp Evans Scuuires [ 238 ] PLATE XLVII EXPLANATION OF THE ILLUSTRATION Prare XLVI. Habit photograph of SoLtanum Torino represented by the collection Schultes 12081 from the Colombian Amazonas. Photograph by Ricuarp Evans Scuurres { 240 | ALVITI PLATE Amazonas, Rio Apaporis, Soratama, between Rio Pacoa and Rio Kana- nari. Altitude about 250 m. ‘‘Flowers greenish. Anthers yellow. Bush. Fruit green, turning orange. Kubeo= be-ta-ka; Taiwano=de- twa; Tatuya=de-twa. Cultivated.’’ September 1, 1951, R. EF. Schultes & 1. Cabrera 13842.—Comisaria del Vaupés, Rio Vaupés, Miti. **Flow- as ers yellow-green. Fruit edible.’’ June 22, 1958, H. Garcia- Barriga, R. E. Schultes §& H. Blohm 15771. Peru: Departamento de Junin, Puerto Yessup. Altitude about 400 m. ° Coarse, erect herb 2-3 ft. Corolla light green, anther yellow. July 10-12, 1929, BF. P. Killip & A.C. Smith 26363,—Departamento del Loreto, Iquitos. Altitude about 100 m. “‘Local name cocona, Coarse herb 3-5 ft. Corolla greenish white. Anthers yellow. Fruit light red, edible. Clearing. ’’ August 2-8, 1929, FE. P. Killip & A.C. Smith 27567. —Departamento del Loreto, Yurimaguas, lower Rio Huallaga, Altitude about 135 m. ‘‘Coarse herb, 3-4 ft. tall. Corolla greenish white. An- thers yellow. Clearing.’? EF. P. Killip & A.C. Smith 27999.—Depart- amento del Loreto, Puerto Arturo, lower Rio Huallaga, below Yuri- maguas. Altitude about 135 m. ““Cocona. Fruit edible, Coarse herb 2-4 ft. Corolla green; fruit red. Clearing.’* August 24-25, 1929, EF. P. Killip & A.C. Smith 67825, Venezur.a: Territorio del Amazonas, Rio Orinoco, San Fernando de Atabapo. 4. Humboldt § Bonpland 918,—State of Bolivar, lower portion of Quebrada Oparu-ma, tributary of Rio Pacairao, below Santa Teresita de Kavanayen. Altitude 915-1065 m. ‘‘Camarata Indian name: bo-pd. In conuco. Herb 2 feet tall. Leaves membranaceous, rich grass-green above, grey-green below. Fruit yellow, edible, made into preserves.”” November 25, 1944, J. Steyermark 60531, Solanum Topiro has never apparently been collected from the wild and, in more than twelve years in the northwest Amazon, I have never seen it outside of agri- cultural plots or abandoned house-sites which obviously had been the scene of cultivation. | believe that we have at hand in this plant a species so long in association with man that it may nowadays exist only because of this association. The fruit yields viable seeds in great abun- dance, but the plants seem to reproduce themselves only in highly disturbed and sunny sites. The Indians eat the ripe fruit as a tomato. The civi- lized inhabitants of the region use the fresh fruits to pre- pare, with sugar, a rather acidulous, thirst-quenching drink. ‘To my knowledge, the plant is never set out de- C242 | PLaTeE XLIX Prare XLIX. Flowering branch of SoLtanum Toprro. Photograph by Ricuarp Evans ScHULTES liberately but springs up from seeds adhering to the rind when this is cast into refuse heaps or when inedible parts of the fruit are spat out in the process of eating. The species is grown over a wide area which includes much of forested eastern Peru, most of the Amazon drainage- area of Colombia, the upper reaches of the Orinoco sys- tem in Venezuela and probably adjacent parts of Brazil. I have never encountered it in Brazil, however, but it may be grown there on a much reduced scale because of the sparsity in the westernmost Amazon of Brazil of unaccul- turated Indian tribes. The type and flavor of the fruit and the heavy bearing characteristics of Solanum Topiro make the plant rather promising as a new subtropical fruit-crop, especially to residents in Florida. As Fennell wrote (loc. cit.): ‘‘As an economic commodity, from the long-range viewpoint, obviously the cocona offers much in the way of improve- ment potential. The complementary values afforded by the various closely related species of this section of the genus are not commonly available to most crop-improve- ment projects. It now remains for us to devise a way to blend and proportion these values satisfactorily toward developing the end product of superior hybrid combina- tions.” In any crop-improvement program, the value of having living material of this rather variable species from all or many parts of its range should not be overlooked, Since Ochse’s notes (loc. cit.) in 1958, apparently noth- ing has appeared which would point to further horticul- tural work in Florida with Solanum Topiro. At that time, Ochse pointed out that nematodes were an im- portant enemy of the plant. The situation is well sum- marized, | believe, by Fennell (loc. cit.) when he states that **. . . a safe assumption is that even in its present unimproved state the cocona is a permanent acquisition as a valuable horticultural plant.” [ 244 | BOTANICAL MUSEUM LEAFLETS HARVARD UNIVERSITY CampripGr, Massacnusetts, JANUARY 16, 1959 Vou 18, No. 6 NEW CONSIDERATIONS IN AN OLD GENUS: DATURA BY ARTHUR S. Barclay THE genus Datura has received considerable attention over the years from ethnobotanists and pharmacologists whose interest in this group of plants stems from the presence of the toxic alkaloids contained in most, if not all, of the species. ‘The herbaceous members of the genus have assumed an outstanding position in the annals of experimental botany at the hands of geneticists and cy- tologists. In spite of the amount of space devoted to this genus in the literature, we are still in need of an adequate taxonomic treatment. It is hoped that this paper will help to clarify some of the nomenclatorial intricacies sur- rounding two species well known to botanists and work- ers in allied fields. Furthermore, descriptions of three new species which were discovered during the preparation of a monograph of the genus are included. There has been much controversy concerning the cor- rect name for the species which has been known as Da- tura meteloides DC. ex Dunal, but which preferably should be called D. inoawia Miller. In order to under- stand how this confusion came about it is necessary to present a brief résumé of the nomenclatorial history of these concepts. [ 245 ] I. Tue Typirication or Datura meteloides DC. ex Dunal The name Datura meteloides was first proposed by Dunal in a manuscript which he sent to Alphonse de Candolle, who in turn published the epithet in his Pro- dromus (1852). Hence, the correct citation should be Datura meteloides DC. ex Dunal. Alphonse de Candolle based his concept of D. meteloides on both the drawing and the description by Sessé and Mocino as we may see from the following note in the Prodromus: ‘‘In calidis Novae Hispaniae regionibus. D. Metel Moe. et Sess. pl. Mexie. ined. ic. et mss. t. 919, collect. transl. Candoll.”’ The original Sessé and Mocino illustrations were loaned to August Pyramus de Candolle and were recalled on short notice. As a result, de Candolle, who recognized the value of these drawings, enlisted the help of the townspeople of Geneva to make copies of them before sending them back. One of these copies, Icones no. 919, has served as the type of D. meteloides and is now pre- served in the Candollean Herbarium (Ewan, 1944). In 1855, a new Datura was brought into cultivation by the French horticulturist M. Louis Vilmorin from seeds sent him by Asa Gray who had obtained them from a collection made by Charles Wright (no. 526) in west- ern Texas in 1849. This plant was cultivated and distrib- uted under the name of Datura meteloides (Kwan, 1944). M. Ortgies, an employee of the Vilmorin Co. of France, noticed the discrepancy between de Candolle’s descrip- tion of D. meteloides and the plant cultivated under the same name and brought it to the attention of the German botanist, Dr. Eduard von Regel. Asa result, a new spe- cies, D. Wrightii, was published by Regel (1859). Gray (1878) considered the southwestern perennial species of the United States to represent Datura mete- [ 246 | loides and relegated D. Wrightii to synonymy handling the obvious lack of agreement between the two descrip- tions with the following comment: ‘*. .. Dunal in DC. Prodr. |.c. 544 (the descr. and drawing of Mocino and Sesse wrong as to 10-dentate corolla).”” This erroneous interpretation was adopted by Safford (1921) who appar- ently followed Gray. It is obvious from Dunal’s monograph (1852) that he had no knowledge of Datura inowia Mill. (Miller, 1786) which, since it is apparently identical with D. meteloides, enjoys priority. Miss Helen Timmerman (1927), although recognizing the problem existing in the literature, failed to reach a satisfactory conclusion regarding D. meteloides and JD. inoxia probably due to insufficient material at hand. Recently, Ewan (1944) became cognizant of the pro- blem concerning the discrepancy between the description of Datura meteloides sensu DC. and our perennial south- western species commonly known by that name. In at- tempting to unravel this confusion he presented an ex- cellent account of the historical backgrounds of both D. meteloides and D. Wrightiit. His conclusion that de Can- dolle’s D. meteloides **cannot be shown to be a misapplied binomial because no other species of Datura has been found in central Mexico which agrees with the original description nor the drawing upon which it is based’’ is hardly acceptable due to the ample herbarium material of D. inoawia at hand. Furthermore, his reeommendation that the name to be applied to our perennial species should be D. Wrightii, provided the identity of D. mete- loides DC. ex Dunal cannot be ascertained, requires more thought. His proposal that the collection C. Wright no. 526 (US 60042) be designated as lectotype of D. Wrightu with Turkey Creek, Uvalde-Kinney Co. line, ‘Texas as type locality appears to be a wise choice, but, [ 247 ] EXPLANATION OF THE ILLUSTRATION Pirate L. The illustration of Datura METELOIDES DC. ex Dunal in the Candollean Herbarium which formerly served as type. [ 248 ] Puatre L 4.3: WA, De 30800 ETT TTT TT[ TY Ty tpt rrr ‘ x Al Ke I Porky sett S44, cm, 1 2 3 4 § rurdutlasntuantariMgant atts tlasssl at a ee ee EXPLANATION OF THE ILLUSTRATION Pare LI. Isotype of Darura MrereLorpes preserved in the Chicago Natural History Museum. Photograph by A.S. Baretay ano R.'T. Moore Prare. LL) Vb) ada! alg spat ils alt ae wiih eed ”. = x/ S47215 HERBARIUM HORT! BOTANIC! MATRITENSIS Piantac Novae Hispaniac. a Seaad, Mocifio, Castillo et Maldonado lectae (I787-1798-1804), Fa eee AAW BEA B= FUSE "Datura Metel N.* EXPLANATION OF THE ILLUSTRATION Prare LIL. Neotype of Darura rnoxta Miller pre- served in the British Museum. Photograph sent by J. BK. Danxpvy PLatE LII CULTIVATED PLANTS Datera. innonia Milley Cultivated at: Chelsea Phogsic Kardon 1843 iL Pe CI PVR MOH 1 et 5 y. A ed ee ir nt Specimen listed on Phil. Trarg £1. 94 no. 'FK3LI 760) aS oma of the 7 thu 9 es . Ede 2 “Fifty Plants From Chelses arden : prsented bo bh Royal Soece ty a the worship ful Com pamy of Ape thecavies for the eae 1757) purtuant to : ee the Direction of Sir Haat Sloume : wT nse according to the present provisions of the International Code of Botanical Nomenclature (Art. 7, Note 8, 1956), C. Wright no. 6.26 should be considered a neotype rather than a lectotype as suggested also by Fosberg in a per- sonal communication to me, Fortunately, | have seen a specimen from the Chicago Natural History Museum which was collected by Sessé and Mocino and labeled by them as **Datura Metel N.” kxcepting for the fruit, this specimen is identical with the illustration of Datura meteloides DC. (see Plates). Because of this outstanding similarity and since there is no other specimen which could fit the needs, | believe that this must represent the type collection. Inasmuch as the Chicago specimen consists of flowering material only, it seems probable that the fruits in the illustration represent an incorrect addition by an unknown. artist. However, I have not yet seen the specimen in the Ma- drid Herbarium (MA). The Sessé and Mocino specimen mentioned above is a duplicate of the original collection. Therefore, the material in the Madrid Herbarium should serve as type specimen, and the Chicago material, Sess, Mocino, Castillo et Maldonado, No. 157.2 ! (KF no. 847215) as an isotype. Datura meteloides, as typified above, is identical with the earlier D. ¢noaia of which it is there- fore, a synonym. It is, furthermore, distinct from JD. Wrightu. Il. Tae TPypreicatrion or Datura inovia Miller Datura inovia was described by Philip Miller (1768) from plants grown from seed which he had received from “La Vera Cruz.”” No type specimen was designated at that time, but there are specimens in the British Museum which were cultivated at the Chelsea Physic Garden dur- ing Miller's curatorship (1722-1770). In the absence of a type specimen, one of these may serve. — 254 ] L The Chelsea Physic Garden, according to Dandy (1958), “‘was conveyed in 1722 to the Society of A pothe- ‘aries by Sloane (who had purchased the manor of Chel- sea in 1712) on certain conditions, one of which was the ‘rendering yearly to the President, Council and Fellows of the Royal Society of London, fifty specimens of dis- tinct plants, well dried and preserved, which grew in their garden the same year, with their names or reputed names; and those presented in each year to be specifi- ‘ally different from every former year until the number of two thousand shall have been delivered’.”” This stip- ulation resulted in the accumulation of an herbarium of over 3000 specimens taken from the Sloane estate. The lists of plants so prepared were printed annually in the Philosophical Transactions (of the Royal Society) from 1723 to 1774. In 1781, the specimens were transferred to the British Museum and are now incorporated in the general herbarium. Miller described the Chelsea plants in successive edi- tions of his Gardener’s Dictionary and as Stearn com- municates, “‘when there is no specimen extant from Miller’s own herbarium. .. . the application of a Miller name can often be determined or checked by a contem- porary specimen from the Chelsea Garden, even though such a specimen was not dried and labelled by Miller himself.” There are, in my possession, excellent photographs of two authentic specimens of Datura inoawia cultivated at the Chelsea Garden and kindly sent by J. fk. Dandy of the British Museum. Having examined these, I hereby designate as neotype the specimen listed in the Philo- sophical Transactions (of the Royal Society) 51 (1760) 99, no. 1845. Since Datura novia and D. meteloides are identical, the former binomial which has priority must be taken up for this concept. ILL. Derscriprions or New SPECIES Datura kymatocarpa Barclay sp. nov. Caulibus dichotomis, usque ad 40 em. altis: foliis in ambitu ovatis, apice acutis, basi inaequalibus, margine irregulariter lobatis vel sinuato-dentatis, usque ad 11 em. longis, dimidio seu duabus-tertiis partibus latis, utrinque strigosis; floribus erectis, axillaribus: pedicellis per an- thesin 0.7-1.8 cm. longis, post anthesin elongatis reflex- isque; calyce cylindrico, 2.8 > 4.6 cm. longo, apice 5- dentato, dentibus triangularibus; corolla albida, infun- dibuliformi, plicata, 6-7.5 em. longa, orificio 5-dentato, dentibus acuminatis, 8-4 mm. longis: capsula globosa, irregulariter dehiscenti, setis semicapillaceis satis dense obtecta, 2.4 cm. longa; seminibus renitormibus, testa verrucosa. Caulescent, dichotomously branching annual herbs, reaching 40 ¢m. or more in height: young branches usu- ally villous, becoming glabrescent with age: leaves alter- nate, ovate in outline with an acute apex and an unequal base extended into a villous up to 8 cm. long petiole: margin irregularly lobate or sinuate-dentate: leaf blades up to 11 em. long and usually one-half to two-thirds as broad, variously strigose on both surfaces except along the mid-vein where it is villous in the same manner as the petiole: pedicellate owers erect, borne in the axils of the branches; pedicels during anthesis 0.7-1.8 em. long, later becoming somewhat elongated and reflexed ; ‘alyx tube cylindrical, 2.8-4.6 em. long and sparsely pu- bescent with a 5-dentate apex, teeth triangular in shape, 3-5 mm. long and 8-5 mm. broad at the base: corolla white, funnel-shaped, plicate, terminating in a 5-dentate orifice, 67.5 em. long: the five corolla teeth acuminate, 3-4.mm. long and 8 mm. wide at the base, each of them supported by three conspicuous nerves, the median one [ 256 ] extending to the tip; the margin between each tooth bisinuate and forming an obtuse lobule giving the corolla the appearance of being 10-angled; the five stamens free, epipetalous, attachedjto the corolla tube at approximately the same level, about 2.5—38 cm. from the base; filaments glabrous, 2-2.3 cm. long; style equalling or slightly ex- ceeding the anthers, 4.6-5.7 cm. long; young ovary cov- ered with spine-like hairs; calyx circumscissile, shed with the corolla, the persistent base expanding into a mem- branous disk-like structure subtending the capsule, 1.5-2 em. in diameter on mature fruit; capsule globose, pubes- cent, pendent on 1.5-2 cm. long pedicels; irregularly dehiscent, covered with semicapillaceous pubescent bris- tles, up to 2.4 em. long; seeds reniform, about 5 mm. long and 3.5 mm. wide, seed coat verrucose. Mexico: State of Mexico, Dist. of Tamascaltepec, Bejucos, ““Ilano,”’ Aug. 15, 1935, G. B. Hinton et al 8173 (Tyrer in US 1841574).—State of Guerrero, Dist. of Coyuca, Pungarabato, ‘‘roadside,’’ July 9, 1935, G. B. Hinton et al 8030 (F, MO, US): Rio Balsas, Aug. 26, 1910, C. R. Orcutt 4384 (F).—State of Michoacan, Municipality of Apatzin- gan, ‘open sun-baked pasture one-half mile S. of Apatzingan, thick adobe soil, alt. 1200 ft.’’, July 31, 1940, W.C. Leavenworth 409 F, GH, MO, NY); Municipality of Apatzingan, “‘common between thorn forest and road, between Apatzingan and La Majada, alt. 1200 ft.°’, Aug. 16, 1941, W.C. Leavenworth & H. Hoogstraal 1615 (F, GH, MO, NY). This species is known only from the Rio Balsas valley in Mexico. According to Leopold (1950), this valley is a tropical inland basin, cut off from the sea by highlands on all sides. In this basin, one encounters a nearly insular type of endemism, and an appreciable number of new species have been described from the collections of Hin- ton and Leavenworth from this general vicinity. This new Datura was collected by Leavenworth in the vicin- ity of Apatzingan at an altitude of 1200 feet and in an area designated by him as open, arid, scrub forest, which [ 257 | covers the greater part of the plains of the valley-floor (Leavenworth, 1946). Datura kymatocarpa may be readily distinguished from all other members of the genus by the semi- ‘apillaceous bristles on its fruit and by its very charac- teristic seeds (see Plates). The specific epithet is derived from the Greek referring to the wavy hairs on the fruit. Datura reburra Barclay sp. nov. Pars quae adest usque ad 80 ecm. alta, caulibus dicho- tomis, glabris; foliis in ambitu ovato-lanceolatis, apice acutis, basi inaequalibus, margine irregulariter lobato- dentatis, usque ad 8 cm. longis, dimidio vel duabus-tertiis partibus latis, lamina utrinque glaberrima; floribus pedi- cellatis, axillaribus, primum plusminusve erectis, deinde post anthesin valde nutantibus; calyce prismatico, cari- nato-angulato, ca. 6cm. longo, apice 5-dentato, dentibus satis prominentibus, ovato-lanceolatis, 1.7—2 cm. longis: corolla infundibuliformi, plicata, usque ad 9.5 em. longa, fauce ampla, 5-dentata, dentibus setiformibus, ca. 5 mm. longis; capsula globosa, puberula, regulariter dehiscenti, pendula, spinis rigidis, acuminatissimis ornata, ca. 2 cm. longa; seminibus reniformibus, lacunis luniformibus utrinque donatis. Dichotomously branching herb at least 80 cm. tall, but surely more (the specimen at hand is incomplete); young branches strigose, becoming glabrous with age, entire plant appearing glabrous except when young; leaves al- ternate, ovate-lanceolate in outline, with an acute apex and an unequal base extended into a puberulent petiole, up to 4.8 cm. in length; margins irregularly lobate- dentate; leaf blades mostly incomplete in our specimen, up to about 8 cm. or more in length, one-half to two- thirds as broad, surfaces essentially glabrous except for the sparsely strigose veins: erect pedicellate flowers borne [ 258 ] in the axils of the branches; the strigose pedicels during anthesis about 1 cm. long becoming elongated and re- flexed with age; prismatic calyx tube 5-toothed at the apex and keeled along the angles, glabrous on both sides, about 6 cm. long; the calyx teeth ovate-lanceolate, 1.7— 2 em. long and 0.5 cm. wide at the base; corolla funnel- shaped, plicate, terminating in a 5-dentate orifice, up to 9.5 em. long; the five corolla teeth setiform, about 5 mm. long and 2 mm. wide at the base, each of them sup- ported by three conspicuous nerves, the median one ex- tending to the tip; exterior of the corolla tube glabrous, the interior pubescent from the base up to the point where the filaments are attached to the corolla tube, then becoming glabrous above this point; the five stamens free, epipetalous, attached to the corolla tube at approx- imately the same level about 3 cm. from the base of the corolla; filaments glabrous except near their attachment to the corolla, 2.8 em. long; anthers about 6 mm. long with filamentous trichomes along the lines of dehiscence; style 7.8 cm. long; calyx circumscissile, forming a re- flexed frill, up to 1.3 cm. long, which subtends the fruit ; capsule globose, about 2 cm. long, regularly dehiscent by four valves, pendent on pedicels up to 2.2 cm. long; fruits covered with stiff, sharply pointed spines; surface of fruit puberulent, spines glabrous; seeds reniform, about 5 mm. long and 4 mm. broad, with lunate lacunae on the lateral faces and with a triple ridge along the con- vex edge which forms a cord-like margin around the seed. Mexico: State of Sinaloa, vicinity of Culican, Sept. 14, 1904, 7.S. Brandegee s.n. (Type: UC 103947). Datura reburra is similar to D. discolor Bernh. in hav- ing both nodding fruit and regularly dehiscing capsules. It is, however, readily distinguishable by its distinctive seeds, its puberulent fruit with glabrous spines and its [ 259 J smaller flowers. The specific epithet of Datura reburra refers to the bristling hairs of the capsule. The question arises, in connection with this new spe- cies, as to the validity of the sections Stramonium Bernh. and Dutra Bernh. The Stramonium-section has been dis- tinguished by erect and regularly dehiscing capsules; the Dutra-section by nodding and irregularly dehiscing cap- sules. Both Datura reburra and D. discolor are charac- terized by having nodding capsules that dehisce regularly by four valves. Thus, they provide intermediates between the two sections in question. Prior to my discovery of Datura reburra, Fosberg had called my attention to the vague distinction between these two sections, pointing out the nonconformity of D. discolor to either one. This additional species would tend to support the belief that these two sections are untenable, unless we propose a new section for every second or third species. Such a procedure would be definitely absurd. In light of the genetic and cytological work on Datura already completed by the late Dr. Blakeslee and by his co-workers, a more thoroughgoing study of these new species should prove to be profitable. Experimental ap- proaches further our knowledge of plants and their rela- tionships to a great degree. Nevertheless, much can also be learned from a study of herbarium material. In fact, such a study should logically precede experimental lines of research. Datura vulcanicola Barclay sp. nov. Planta fruticosa, usque ad 8 m. alta; foliis ovatis vel ovato-ellipticis, apice acutis, basi in petiolum decurrenti- bus, utrinque puberulescentibus, margine integra vel irregulariter sinuato-dentata; floribus solitariis, plus- minusve horizontaliter cernuis vel nutantibus arcuatis; calyce glabro, paulo inflato, cylindrico, 4.5—8 cm. longo, [ 260 ] apice inaequaliter 2-4 dentato; corolla usque ad 20 cm. longa, urceolata, apice 5-dentata, margine inter dentes breviter retusa; capsula obovata, prominenter bisulcata, irregulariter verrucosa, usque ad 12 cm. longa; semine reniformi, testa glabra; numerus chromosomicus, 2n= 24, Perennial shrub or small tree reaching 8 meters or more in height; leaves alternate; blades up to 20 cm. long and up to 10 em. broad, puberulent on both surfaces, ovate to ovate-elliptic in outline with an acute apex and a sub- equal base decurrent on the up-to-10 cm. long petiole; margins entire to irregularly sinuate-dentate; the larger, dentate leaves occurring on juvenile shoots with the smaller and entire ones on older branches; flowers axil- lary on somewhat arcuate, 1.5-2.7 cm. long pedicels; corolla tube basally green becoming light red which fades into yellow near the mouth, the veins red, interior of the tube deeply red tinged between the nerves, the upper half yellowish white, becoming deeply yellow near the mouth; calyx glabrous, somewhat inflated, cylindric, 4,.5-8 cm. long, often split along one side giving a spathe- like appearance to it; calyx teeth two to four, unequal in length; corolla up to 20 cm. long, urceolate, terminat- ing in a 5-dentate orifice, each tooth supported by three prominent nerves; the margin between the short and acuminate teeth slightly retuse; the five stamens free, epipetalous, extending to just below the mouth of the corolla, attached to the corolla tube at approximately the same level, 7-8.9 cm. from the base; filaments gla- brous above, becoming villous near the point of adnation, anthers white, filaments pale green; shortly after anthe- sis the calyx dehisces at the base of the ovary and is shed; capsules obovate, deeply bisuleate with a secondary shal- low division, up to 12 cm. or more in length, warty in appearance; seeds reniform, about 7 mm. long and 5 mm. wide, seed coat smooth on both sides, shiny, slightly [ 261 J EXPLANATION OF THE ILLUSTRATION Pirate LIII. Upper: Fruit of the type of Datura reBURRA Barclay (TS. Brandegee, s.n., Sept. 14, 1904). Lower: Fruit of the type of Darura KyMa- rocarpa Barclay (G. B. Hinton, No. 8173). Photograph by Cuaries DILts [ 262 ] PLaTE LIII | 1 | | HLT EXPLANATION OF THE ILLUSTRATION Prare LIV. Upper: Seeds of Datura KyYMATOCARPA Barclay (Leavenworth, No. 1615). Lower: Seeds of the type of Darura resurra Barclay (T. 8. Brandegee, s.n., Sept. 14, 1904). Photograph by Cuarces Ditties [ 264 | » LIV PLATE EXPLANATION OF THE ILLUSTRATION Prare LV. Darura vetcantcoLta Barclay. Photo- graph of the plant from which the type specimen was collected (4.8. Barclay & R. E. Schultes No. 147). Photograph by Ricuarp Evans ScHULTES [ 266 ] EXPLANATION OF THE ILLUSTRATION Prare LVI. Darura vutcantcoia Barclay. Same as Plate LV in flower and fruit. Photograph by Ricuarbd Evans Scuurres [ 268 ] LVI PLATE ridged on the convex side; chromosome number, 2n= 24. Cotompra: Dept. of Cauca, Municipality of Puracé, northern slope of the Volean de Puracé, alt. 2700-2800 m., July 28, 1956, 4.8. Barclay & R. FE. Schultes 147 (Vyvn: GH).-——Same locality and date, A. S. Barclay & R. EF. Schultes 149 and 177 (GH).—Between Puracé and its paramo, Chiquin, heath-brush, alt. 2700-3100 m., July 11, 1989, E. Pérez-Arbeldez & J. Cuatrecasas 5960 (F, US, COL.).—Pu- racé, alt. 8800 m., Feb. 1938, A. von Sneidern 1S9S (US).—Paletara to Calaguala, alt. 83000-8200 m., July 17, 1922, F. WW. Pennell 70928 (GH, NY, US). In 1956, during my field studies of the tree Daturas in the Colombian Andes, | was surprised to find on the side of the famous and active Volcan de Puracé popula- tions of what was at once distinguishable as a striking and beautiful new species of the genus. Herbarium speci- mens, fixed buds tor cytological study, about 50 pounds of leaves tor chemical analysis and living material in the form of cuttings and seeds for horticulture were gathered. Karlier specimens which had been collected in the same general vicinity on the slopes of the volcano and which also represent the new species had been filed in our her- baria under the name Datura sanguinea Ruiz & Pavon. D. vuleanicola seems to be most closely related to J. sanguinea on the Basis of comparative morphology. Although Datura sanguinea is a highly variable spe- cies, D. vuleanicola may readily be distinguished from it by a number of characters, among the most prominent of which are its warty fruit and its smooth seeds. The fruit has a hard, almost woody pericarp and is attached to the branch by a stout peduncle which becomes nearly as thick as the branch supporting it. J. sanguinea, on the other hand, is characterized by its smooth, more fleshy fruit which is attached by an elongated, relatively un- thickened peduncle and by its verruculose seeds. The flowers of the new species are cernuous or nodding at a 80 to 45 degree angle, with a glabrous, nearly satin- textured calyx and corolla. The light red to salmon- colored corolla tube fades into yellow near the mouth. The nerves are red, and the slightly inflated calyx tube dehisces after anthesis. In comparison, the flowers of Datura sanguinea are vertically pendulous, variously pubescent, and although the coloration is quite variable, the corolla tube is usually yellow becoming red towards the mouth with conspicuous greenish to yellow nerves. A much inflated calyx tube which is often persistent following anthesis forms a husk around the fruit. Datura vulcanicola, as far as | could judge, does not reach the great height or corpulence which is commonly found in D. sanguinea. The chromosome number of 2n =24 (4. S. Barclay & R. FE. Schultes no. 147) agrees with that of the other arborescent Daturas investigated. In a communication to me, Professor Schermerhorn of the Massachusetts College of Pharmacy has related that a preliminary biochemical analysis of the leaves of this new species has shown no significant differences be- tween the alkaloids present in Datura vulecanicola and in the other species of Datura. Although Datura vulcanicola occurs in large numbers in the subpdiramo heath vegetation under conditions which could be interpreted as wild, | am of the opinion that the abundance of individuals at the type locality is probably the result of human activity. My principle reason for believing this is that most of the plants were growing along what are or obviously had been foot paths or donkey trails. Furthermore, we must not overlook the fact that for centuries this agriculturally rich volcanic area has been populated by advanced Indian peoples who are known to have employed Daturas as narcotics in their magic and therapeutic practices. It is interesting to note in this connection, however, that 2. vulcanicola has not been spread, at least to any appreciable extent, from this | 271. | locality by primitive man. It may yet be found on neigh- boring mountains, but neither the comparatively thor- ough explorations in this general area in the past nor my own collecting, directed almost exclusively to Datura, have turned it up elsewhere. ACKNOWLEDGMENTS | would like to acknowledge the help and encourage- ment of my major professor, Dr. R. C. Rollins, who made available to me the Fernald Fund of Harvard Uni- versity for field work in South America. I also owe a debt of gratitude to Dr. R. E. Schultes who first sug- gested the genus Datura asa research subject and whom I had the privilege of accompanying while in South Amer- ica. To Dr. Jesis M. Idrobo, Dr. Alvaro Fernandez- Pérez and the other members of the Instituto de Ciencias Naturales, Bogota, Colombia, I wish to extend my deep- est gratitude and appreciation for their kindness and help during my stay in their country. BIBLIOGRAPHY Dandy, J. E. 1958. The Sloane Herbarium. British Museum, Lon- don. 166. Dunal, F. 1852. Solanaceae [Datura] in Alphonse de Candolle Pro- dromus, 13, pt. 1: 538-546, Ewan, J. 1944. Taxonomic history of the perennial southwestern Datura meteloides. Rhodora 46: 317-328. Gray, A. 1878. Synoptical Flora of North America, 2, pt. 1: 240. Leavenworth, W. C, 1946. A preliminary study of the vegetation of the region between Cerro Tancitaro and the Rio Tepalcatepec, Michoacan, Mexico. American Midland Naturalist. 36: 137-206. Leopold, A. Starker. 1950. Vegetation zones of Mexico. Ecology. 31: 507-518. Miller, P. 1768. The Gardener’s Dictionary, ed. 8. (Datura n. 5). Regel, KE. von. 1859. Datura Wrightii Hort. Gartenflora 8: 198, t. 260. Safford, W. E. 1921. Synopsis of the genus Datura. Jour. Wash. Acad. Sci. 11: 173-189. Timmerman, H. 1927. Datura: the nomenclature of the species used in medicine. Pharm. Jour. (London) 118: 571-574. [ 272 ] THE PRESENT DISTRIBUTION OF NARCOTICS AND STIMULANTS AMONGST THE INDIAN TRIBES OF COLOMBIA BY Nesror Uscarecut M.’ RECENT extensive and intensive interest in narcotic and stimulant plants on the part of medical and pharmacolo- gical investigators has focused attention on the need of knowing more about these plants from the botanical and anthropological points of view. During the past twenty years or so, significant strides have been made in deter- mining the correct botanical sources of plant narcotics and stimulants. Very little has been done, however, to- ward bringing into some semblance of order the vast amount of information on their distribution, preparation, use and social significance, which is scattered here and there throughout ethnological literature and the writings of travellers. Forasmuch as Colombia represents one of the regions of the world where the native population has developed to its highest degree the use of plants which act on the central nervous system as intoxicants or stimulants, I propose here to present a preliminary and very much abbreviated survey of the distribution of the use of these plants amongst the Indians of this country. "Instituto Colombiano de Antropologia, Bogota, Colombia. [ 278 | L I. The population of Colombia is a racial mixture that is constantly increasing in complexity. It consists in great part of Europeans, especially from Spain, and of Indians from Middle America, the Antilles, the northern Andes and the Amazonian regions. There are also some ne- groes, mostly from Africa, who have intermingled with both whites and Indians, especially in some of the De- partments. All these groups have contributed their share to the spiritual and cultural complex which man repre- sents in Colombia. Much of Colombia, sparsely populated and with a dearth of roads, is occupied by Indian tribes which sub- sist wholly on local resources. The rivers provide these natives with roads, meat and drink; the plant kingdom furnishes their clothing, shelter, food, medicines, stimu- lants, narcotics and poisons; the animal world supplies food, diversion and other material for magic. These Indians employ sundry stimulants and narcotics of great potency in both magico-religious rites and in daily life as well. The accompanying map shows the wide distribution of the use of such plants. This is so great that it causes us to wonder why such a complex exists in Colombia. Many of the naturalists, ethnographers, explorers and geographers who have lived or travelled amongst these Indians have attempted to interpret the reason for the extensive use, and even near-abuse, of stimulants and narcotics in this area. Actually the reason is simple: it is due merely to primitive mentality which is mystical. The Indian considers all nature and the visible and im- agined cosmos to be endowed with spirit-forces. He does not recognize any boundary between natural and super- natural phenomena. This reason is in concord with that of Pardal (27) and Frazer (14), but is not in agreement [ 274 | Piatt LVII a ae eae aa - SS ee ere ee SgQ&™ ON Py I cere SLA L SA \ eer a “Ro: — Za oss aR Texas Agric. Exp. Sta. Bull. 730: 1-18. ~ . . P oe Rosbaco, U. F., 1951. Consideraciones sobre maices amargos’’ con especial referencia a su cultivo en la provincia de Entre Rios. Idia No. 46: 1-12. . Weatherwax, P., 1935. The phylogeny of Zea Mays. Amer. Midland Nat. 16: 1-71. ——, 1950. The history of corn. Sci. Month. 71: 50-60. [ 410 | 53. 57. Wellhausen, E. J., L. M. Roberts and E. Hernandez X. in collabo- ration with P. C. Mangelsdorf, 1952. Races of maize in Mexico. Bussey Institution, Harvard Univ. 1-223. —— and C, Prywer, 1954. Relationship between chromosome knob number and yield in corn. Agron. Jour. 46: 507-511. . ——, A. Fuentes O. and A. Hernandez C., in collaboration with P. C. Mangelsdorf, 1957. Races of maize in Central America. Nat. Acad. Sci.—Nat. Res. Council Publ. 511: 1-128. Vachhani, M. V., 1950. A study of the relationship of chromo- some knobs with certain agronomic and morphological characters in corn inbreds. Agron. Jour, 42: 196-201. Venkatraman, T. S. and R. Thomas, 1932. Sugareane-sorghum hybrids. Indian Jour. Agric. 2: 19-27. [ 411 ] BOTANICAL MUSEUM LEAFLETS HARVARD UNIVERSITY CamsripGak, Massacuusetts, Apri, 17, 1959 VoL. 18, No. 10 THE ORIGIN OF CORN ITV. PLacre AnD TIME oF ORIGIN BY Pau. C. MANGELSDORF AND Ropert G. REEVES In regard to the origin of this plant, although there has never been room for reasonable doubt, there have been those who fancied there was room for argument. America is clearly and beyond question the native country of Indian corn. Yet, from the commencement of its history, writers have not been wanting to contest this point, and to claim for it an Eastern origin. The weight of authority and of argu- ment so entirely preponderates in favor of its American origin, that it is searcely worthwhile, in a work aiming to be useful rather than learned, to waste the time of the reader with idle and unprofitable speculation, Epwarp Enrietp (10). This statement which appeared almost a century ago in an otherwise undistinguished work is as true today as it was then. We supposed when, some twenty years ago, we wrote our monograph on the origin of Indian corn and its relatives (81) that there was one question — its place of origin, America or the Old World — which had been answered once and forall. It turns out that we were wrong; for although our monograph stimulated much useful interest in the problem of the origin of maize, it also opened a veritable Pandora’s box of unrestrained speculation on certain aspects of the problem. It has, as a consequence, become necessary once again to review the evidence pertaining to the question of corn’s place of origin. [ 413 ] PreE-CoLUMBIAN MAIZE IN Asta? Among the herbalists and early botanists who gave their attention to maize, there were a number who re- garded it as a plant of Old World origin (cf. 30). The evidence which de Candolle (7) marshalled to support his conclusion on the American origin of maize was so con- vincing that the problem was then generally regarded as solved. But several times in this century the question of an Asiatic origin or of a pre-Columbian distribution of maize in Asia has been raised. Following the discovery of a previously unknown type of endosperm, ‘“‘waxy,”’ in a variety of Chinese maize, Collins (9) suggested, de- spite Laufer’s (23) earlier conclusion to the contrary, that maize may have been known in Asia before the discovery of America. More recently Anderson has, on several occasions (1, 2), suggested the possibility of an Asiatic origin of maize or of its prehistoric spread to Asia, and, in a joint paper with Stonor (38) describing some collec- tions of maize from Assam, reached the conclusion that ‘‘maize must either have originated in Asia or have been taken there in pre-Columbian times.” The conclusions of Stonor and Anderson were wel- comed by the ‘‘diffusionists,’* a school of geographers, anthropologists and others who professed to see in art forms, myths, and other cultural traits, including the use of plants, great similarities between Asia and America and who, for reasons which are not at all clear to us, are apparently determined to prove that all of these traits diffused from a common center. So far as plants are con- cerned the diffusionists’ theses have had the support, especially of Carter (8), Heyerdahl (15), and Sauer (37), all of whom have regarded the conclusions of Anderson or of Stonor and Anderson as supporting the idea of pre- Columbian, trans-Pacific diffusion. [ 414 ] Diffusionists may also have found encouragement in Hatt’s (14) fascinating study of American and Indone- sian folklore which showed some remarkable similarities between the two, especially with respect to various ver- sions of the Corn Mother myth. For example, the origin of cultivated plants from a sacrificed child, an important motif in Indonesia, is also conspicuous in Peruvian myth- ology. Nevertheless, Hatt was compelled to conclude that if agriculture and myths reached America across the Pacific Ocean this must have taken place not all at once but at different times, He did not suggest diffusion in the opposite direction nor did he consider it a possibility, as Sauer seems to have, that a Corn Mother myth could have been diffused except as it accompanied the spread of a grain. The idea of a pre-Columbian interchange of plants be- tween the Old World and the New was virtually de- molished by Merrill (83). He showed that there is not only a lack of tangible evidence for such prehistoric dif- fusion but also that the presence of American plants in Asia soon after the discovery of America is easily and reasonably accounted for by the early Portuguese trade route established in 1500 from Brazil to Goa by way of the Cape of Good Hope. So far as maize is concerned, the case for its pre- Columbian occurrence in Asia, never a very convincing one, was considerably weakened when Mangelsdorf and Oliver (30) showed that the Assamese maize described by Stonor and Anderson is not at all unique and has close counterparts in Colombia and other parts of South America. The case has recently been weakened still more by new evidence presented by Ho (16) who, after a searching study of Chinese historical sources, concluded that maize was introduced into China early in the six- teenth century arriving there by both overland and mari- time routes. He states: [ 415 ] Summing up the introduction of maize into China, we may say that maize was introduced into China two or three decades before 1550; that it was probably introduced by both the overland and maritime routes ; that there is little reason to justify Laufer’s far-reaching conclusion, especially in the light of the introduction of other New World plants, that in the dissemination of food plants ‘a land route is preferred over a sea route as their way of propagation’; and that, barring a sensa- tional discovery in Chinese sources clearly indicating a pre-Columbian introduction, Chinese maize as a topic for anthropological speculation should be closed. Sut6 and Yoshida (39) were doubtless unaware of Ho’s paper when they concluded, on the basis of decid- edly meager evidence, that one of the types of oriental maize, Persian, described by them must have had an ex- tensive pre-Columbian distribution in parts of Asia. They also favored Anderson’s suggestion (2) that corn originated in Asia perhaps as an amphidiploid of a five- chromosome species of Coiv or Sorghum. They were apparently unaware also of the discovery by Barghoorn et al (4) of fossil maize pollen in the Valley of Mexico almost identical with that of modern maize pollen. This discovery, to be discussed in more detail later, virtually proves the American origin of corn and rules out the pos- sibility of an Asiatic origin. PRE-COLUMBIAN CORN IN AFRICA / The confusion which can result from what Enfield has called ‘‘idle and unprofitable speculation’’ is nowhere better illustrated than in Jeffreys’ (20) acceptance of that part of the Stonor-Anderson thesis which holds that, if maize did not originate in Asia, it must have been taken there in prehistoric times. Jeffreys (17) had earlier assembled extensive historical references purporting to show that there had been Arab- Negro contacts with the Americas beginning about 900 A.D. and that maize had been introduced into Africa before 1492. When Goodwin (12) described potsherds [ 416 | from Ife in Nigeria, apparently decorated by rolling a maize cob over wet clay, Jeffreys (18) proceeded to date the introduction of maize into the region in Africa repre- sented by Ife at 1000-1100 A.D.', a date slightly earlier than the one which he had arrived at on the basis of other evidence (19). He then showed by linguistic and historical evidence how it might have spread from Africa to Asia (20). To analyze Jeffreys’ arguments in detail would seem to serve no useful purpose until it should first become clear: (A) that the impressions on the African pottery are unmistakably those of maize*; (B) that they are un- mistakably pre-Columbian. Unless these two facts can be clearly established, we prefer to agree with Goodwin’s recent statement (in a letter) which he has kindly given us permission to quote: . and am of the opinion that not all of this pottery was decorated by rolling a maize cob over the surface. I have no evidence from that or from any other source suggesting that maize reached Africa in Pre- Columbian times. PRE-COLUMBIAN MaIzE IN Europe? Finan (11), ina study of the maize illustrated and de- scribed in the herbals, concluded that there were two ' Weatherwax erroneously attributes to Goodwin the idea of a pre- Columbian introduction of maize into Africa. Goodwin carefully avoided drawing such a conclusion, ? We have not been able to obtain specimens of the African pot- sherds for examination but, since this was written, we have seen photographs of one of them displayed at the Tenth International Con- gress of Genetics in Montreal by Dr. W.R. Stanton of Nigeria. There is little doubt that this impression is of a maize cob since the paired arrangement of the spikelets is clearly shown. But Stanton, like Goodwin, regards these impressions as post-Columbian and states that he is in general agreement with Porteres (35) who, after carefully re- viewing the evidence presented by Jeffreys, Mauny, and others, con- cluded that maize reached Africa in the sixteenth century by two routes: a flint corn by way of the Mediterranean and the Nile; a soft [probably dent] corn by way of the coast of Guinea. [ 417 ] distinct types: the first, characterized by conspicuous prop roots, was probably a tropical form introduced into Europe from the Caribbean area soon after 1492; the second, which lacks prop roots but sometimes has nu- merous tillers, is similar to the Northern Flints of east- ern North America and appears to have been well known in Kurope within 50 years after America’s discovery. He (as well as Anderson in the preface to Finan’s work) raised the question whether it could have been intro- duced into Europe by the Norsemen before 1492. Finan also speculated on the reason for the common. belief among the herbalists that corn came to Europe from the Orient, Suto and Yoshida have gone even further than this in their unqualified assertion that the Aegean type, from which they believed the European maize to be derived and which was first described by Anderson and Brown (3), was, like the Persian, diffused throughout the Old World before 1492. Corn’s Rapip SPREAD AFTER 1492 If all of these various assertions about pre-Columbian maize inthe Old World were true, maize must have been about as widely distributed there as it was in America. Why then did it not leave a single tangible record of any kind of its presence? Why did corn cause such wonder- ment to sixteenth century students of plants if it had already been known for several centuries or more? Underlying all of the speculations on pre-Columbian maize in Asia, Africa, or Europe is one common assump- tion: that corn could not have spread rapidly enough after 1492 to reach all of the places where it was known a generation later. This is not only a highly unreliable premise but also, we think, a presumptuous one for it places arbitrary limits, not justified either by history or [ 418 | by contemporary experience, on mankind’s capacity to spread, through trade and other means, the world’s prod- ucts. So far as Europe and Africa are concerned, the early post-Columbian occurrence of maize is explained quite satisfactorily by Wright (46) who showed how the Moors, after being partially expelled from Spain between 1499 and 1502, took maize with them to Tangier and the north African coast whence it rapidly spread to that part of the world which lay around the Mediterranean Sea, i.e., Turkey, Syria, and Egypt. Wright explains further that the name ““Turk’’ in England during the sixteenth century was often used indiscriminately with ‘‘Moor”’ to indicate a Moslem. It seems probable, therefore, that maize at one period was obtained more easily in western Europe from the Moslem regions of the Mediterranean than from the West Indies and hence was known to the English as ‘“Turkey corn’’ (both Egypt and Syria were then parts of Turkey) and to the Italians as grano turco. Perhaps the belief, held by a number of the herbalists, that maize came from the East was based on nothing more than the fact that its common name seemed clearly to indicate an eastern origin. Some recent modern con- clusions regarding its origin have had little more foun- dation in fact. THE PLACE oF ORIGIN IN AMERICA The discovery by Barghoorn et a/ (4) of fossil pollen in the Valley of Mexico seems now to have established the origin of corn in America beyond question but it still leaves open the problem of where in America maize was first domesticated. The fossil pollen also proves without doubt that wild maize once grew in the Valley of Mexico. But the fact that maize pollen was found in the drill core only at great depths (below 69 meters) and then was absent until it [ 419 ] appeared again at the upper levels (above 8.6 meters), probably after the establishment of agriculture, suggests that the early fossil maize was that of a colony which became extinguished, perhaps through volcanic action. However, if wild maize grew in one Mexican valley it may well have grown in others and in similar sites in other regions. Archaeological maize from caves in Mexico and New Mexico—some of it not far removed in its characteristics from wild corn—also points to an early center of domes- tication in Mexico. Furthermore, the oldest archaeolo- gical corn so far discovered in South America—that de- scribed by Bird (5) from Huaca Prieta—is later than the earliest corn from either Bat Cave or La Perra Cave and is more advanced in its development. Finally, anthro- pologists now tend to believe that the prehistoric cultures of America had their beginnings in Middle America and spread from there to South America (45). For all of these reasons, Mangelsdorf concluded sev- eral years ago (26) that ‘‘Maize undoubtedly had at least one center of origin in Middle America.” This conclusion is, of course, directly contrary to our earlier one (31) that maize had its origin in the lowlands of South America—an assumption based on the fact that pod corn, which we then regarded and still regard as the ancestral form, was repeatedly encountered there and was apparently unknown in Mexico and Central Amer- ica. And then, too, with teosinte disposed of as a hybrid of maize and Tripsacum, there no longer seemed to be any compelling reason for looking to the region where teosinte is native as the center of origin of maize. We have never been convinced, as Weatherwax has appar- ently been (41, 42, 43), that corn’s center of origin must coincide with the center of diversity of its relatives, teo- sinte and ‘Tripsacum. [ 420 | In spite of the recently discovered evidence for a Mid- dle American origin of cultivated maize—one which has been favored on the basis of other evidence not only by Weatherwax but also by Kempton and Popenoe (21), Kuleshov (22), Meade (82), and Vavilov (40)—we are not yet ready to rule out completely the possibility of an independent center of origin somewhere in South America although our earlier idea of a single origin in the lowlands of South America has now been abandoned. There are still too many facts which are not completely explained by the assumption of a single origin in Middle America. Among these are: (a) the great diversity of corn in the highlands of Peru; (b) the fact that all of the known pericarp colors of corn occur in one depart- ment, Ancash, of Peru (13); (c) the frequent occurrence of pod corn in valleys on the eastern slopes of the Andes; (d) the high incidence of the tu” gene in Peruvian corn (27); (e) the occurrence in Peru of a primitive race, Con- fite Morocho, which could conceivably be the progenitor of all of the other known primitive races of the hemi- sphere (18). So far as the evidence from living corn vari- eties is concerned, it still points strongly to a South American center in the highlands of Peru, Bolivia and Ecuador and, were it not for the conflicting evidence from fossil pollen and archaeological maize, we should unhesitatingly continue to assume that corn had its ori- gin in South America. An obvious solution to this dilemma is to assume that maize has been domesticated more than once. Such an assumption would not be radically new. We (81) have previously pointed out that five of the cultivated plants common to Middle America and South America— squashes, beans, tomatoes, amaranths, and cotton—were represented in the two regions by different species or sub- species. There is now evidence that this may have been [ 421 ] true also of corn. There are at least three different prim- itive races of maize in Peru today (13); two in Colombia (36); and four in Mexico (44). Precursors of two of the Mexican races, Nal-el and Chapalote, which in their characteristics are not far removed from wild corn, have been identified in archaeological collections from caves in northeastern Mexico (29) and northwestern Mexico (28). This shows that even in quite early stages of domestica- tion there were already distinct types of corn. One con- clusion which might be drawn at this time is that wild corn occurred sporadically in restricted sites in the moun- tainous region of this hemisphere: in Bolivia, Ecuador, and Peru in South America and in Guatemala and Mex- ico in Middle America. Once agriculture had been in- vented and maize domesticated it may have been domes- ticated repeatedly by the American Indians wherever it was found. Sucha conclusion might require modification at any time as the result of the discovery of new archae- ological evidence, especially from South America.” In this connection mention should be made of the hy- pothesis of Birket-Smith (6), based largely on linguistic evidence, that maize originated in Colombia, perhaps in the lower Rio Magdalena Valley. In Colombia there are 3 Since this was written a study (still unpublished) of the most re- cent find of archaeological maize from a site in the Ica Valley on the coast of Peru shows that the predominating type of corn is remark- ably uniform and is similar, if not identical, to the prehistoric precur- sor of the still existing Mexican race, Chapalote. Some of the Ica ears, however, show various modifications which can be attributed to introgression from a race of popcorn, Confite Morocho, which is still found in parts of Peru, especially in the Department of Ayacucho. It now appears that the great diversity of maize in Peru had its begin- nings when a prehistoric popcorn from Mexico hybridized with the Peruvian popcorn, Whether the Peruvian popcorn was already in cul- tivation when the Mexican race was introduced cannot be determined from the evidence now available. Additional archaeological evidence from the Peruvian highlands may shed new light on this problem. [ 422 ] ‘ names, some of which are regarded as ‘‘primitive,’’ as- sociated with maize, its culture and uses, which have affinities with those of Central America, Ecuador, Peru, Venezuela, and the South American lowlands. Roberts et al (36) have suggested that such a situation might have developed if this region had been not a center of origin but a crossroads in which the cultures of Central Amer- ica, the Andean highlands, and the South American lowlands converged. That northern Colombia was defi- nitely a crossroads region is now generally accepted by anthropologists. Evidence from a study of Colombian races of maize tends to support this interpretation. Other aspects of the case for a Colombian origin have been re- viewed by Mesa Bernal (34). THe TIME or ORIGIN The fossil maize pollen of Mexico, presumably that of a wild corn, is assigned, on the basis of systematic changes in the frequency of other types of associated pollen, to the last interglacial period. Recent estimates place this at 80,000 years or more ago. There is no reason to doubt that wild maize is much older than this. The origin of cultivated maize is, of course, much more recent. The oldest archaeological specimens so far stud- ied, those of Bat Cave in New Mexico, are dated by Libby’s radiocarbon determinations of associated char- coal at 5,600 years. There is a possibility that the pre- historic maize and charcoal are not contemporaneous; that the charcoal is a residue of fires built by itinerate campers long before the cave was occupied by maize- growing people. ‘This date, however, is not inconsistent with those from other sites. The oldest corn from La Perra Cave, dated by radiocarbon determination of asso- ciated vegetal remains at 4445 years, is by no means as primitive as the earliest Bat Cave specimens. The oldest [ 423 ] archaeological maize from Swallow Cave, similar to the earliest Bat Cave material, has not been dated but the fact that it occurred in levels 13 and 14, seven feet below the surface, suggests a very substantial age. The pre- pottery corn from Tm C 247, a site excavated by MacNeish (24, 25), some of which is similar to the Bat Cave corn, has been tentatively dated at 8945 years. The oldest corn from Huaca Prietain Peru, dated about 2900 years, is already well advanced in its development over the earliest Bat Cave corn. The evidence, so far as it goes, is consistent with the conclusion that corn was first domesticated about 5000 years ago or perhaps a millennium or more still earlier. How, even in this length of time, could the primitive corn, with which domestication began, have evolved into the highly developed varieties of today such as the Corn- Belt corn of the United States with its magnificent ears or the spectacular large-seeded flour corn of the region of Cuzco, Peru? This is a question to which we hope that the earlier papers in this series will have given, at least, some of the answers. [ 424. ] as 10. 1H 12, iS. 14. LITERATURE CITED . Anderson, E., 1943. A variety of maize from the Rio Loa. Ann. Missouri Bot. Gard. 30: 469-474, ——, 1945. What is Zea Mays? Chronica Botanica 9: 88-92. —— and W.L. Brown, 1953. The popcorns of Turkey. Ann. Missouri Bot. Gard. 40: 33-48. Barghoorn, E. S., M. K. Wolfe and K. H. Clisby, 1954. Fossil maize from the Valley of Mexico. Bot. Mus. Leafl. Harvard Univ. 16: 229-240, Bird, J., 1948. America’s oldest farmers. Nat. Hist. 57: 296- 303, 334-335. Birket-Smith, K., 1943. The origin of maize cultivation. Kel. Danske Videnskabernes Selskab; Hist.—Filol. Meddel 29: 1- 49, . Candolle, A. de, 1886. Origin of cultivated plants. Reprint of 2nd edition, 1959. New York, Hafner Publishing Co. . Carter, G. F., 1950. Plant evidence for early contacts with Amer- ica. Southwest Jour. Anthrop. 6: 161-182. Collins, G. N., 1909. A new type of Indian corn from China. U.S. Dept. Agric. Bur. Plant Industry Bull. 161: 1-30. Enfield, E., 1866. Indian corn, its value, culture and uses. New York, D. Appleton and Co. Finan, J. J., 1950. Maize in the great herbals. Waltham, Mass., Chronica Botanica. Goodwin, A. J. H., 1953. The origin of maize. S. African Arch. Bull. 29: 13-14. Grobman, A., W. Salhuana and P. C. Mangelsdorf, 1956. Races of maize in Peru. Maize Gen. Codp. News Letter 30: 27-30. Hatt, G., 1951. The Corn Mother in America and in Indonesia. Anthropos 46: 853-914, Heyerdahl, T., 1952. American Indians in the Pacific; the the- ory behind the Kon-Tiki expedition. London, George Allen & Unwin Ltd. 18. 20. 21. fas) a) 24, 26. - 28. 30. Ho, P. T., 1956. The introduction of American food plants into China. Amer. Anthrop. 57: 191-201. . Jeffreys, M. D. W., 1953. Pre-Columbian negroes in America. Scientia (July-Aug.), 1-18. ——, 19538. Pre-Columbian maize in Africa. Nature 172: 965- 966. . ——, 1958. The history of maize in Africa. Eastern Anthrop. 7: 138-147. ——, 1955. Pre-Columbian maize in Asia, Eastern Anthrop, 9: 21-28. Kempton, J. H. and W. Popenoe, 1937. Teosinte in Guatemala. Carnegie Inst. Washington Publ. 483: 199-218. Kuleshov, N. N., 1929. Geographical distribution of the diverse varieties of maize in the world. Bull. Appl. Bot. Gen. Pl. Breed- ing 20: 506-510. Laufer, B., 1907. The introduction of maize into Eastern Asia, Proce. 15th Intern. Congr. Americanists 1: 223-257. MacNeish, R. S., 1955. Ancient maize and Mexico. Archaeology 8: 108-115. . ——, 1958. Preliminary archaeological investigations in the Sierra de Tamaulipas, Mexico. Trans. Amer. Philos. Soc. 48, pt. 6: 1-210. Mangelsdorf, P. C., 1954. New evidence on the origin and an- cestry of maize. Amer. Antiquity 19: 409-410. ——, 1957. Half-tunicate from Peru, Ecuador and Paraguay. Maize Gen. Codp. News Letter 31: 64. and R. H. Lister, 1956. Archaeological evidence on the evolution of maize in northwestern Mexico. Bot. Mus. Leafl. Har- vard Univ. 17: 151-178. , R. S. MacNeish and W.C. Galinat, 1956. Archaeological evidence on the diffusion and evolution of maize in northeastern Mexico. Bot. Mus. Leafl. Harvard Univ. 17: 125-150. and D. L. Oliver, 1951. Whence came maize to Asia? Bot. Mus. Leafl. Harvard Univ. 14: 263-291. [ 426 ] 31. 32, 36. Si. 38. 39. 40. 41. 42. 43, 44, 45, 46. Mangelsdorf, P. C. and R. G. Reeves, 1939. The origin of Indian corn and its relatives. Texas Agric. Exp. Sta. Bull. 574. Meade, J., 1948. Iziz Centli (El Maiz). Mexico. . Merrill, E. D., 1954. The botany of Cook’s voyages. Waltham, Mass., Chronica Botanica. Mesa Bernal, D., 1957. Historia natural del maiz. Revista Acad. Colomb. Cien. Exact. Fis. Nat. 10: 13-106. Porteres, R., 1955. L’introduction du mais en Afrique. Jour. Agric. Trop. Bot. Appl. 2: 221-231. Roberts, L. M., U. J. Grant, R. Ramirez E., W. H. Hatheway and D. L. Smith, in collaboration with P. C. Mangelsdorf, 1957. Races of maize in Colombia. Nat. Acad. Sci.-Nat. Res. Council Publ. No. 510. Sauer, C.O., 1952. Agricultural origins and dispersals. New York, Amer. Geog. Soc. Stonor, C. R. and E. Anderson. 1949. Maize among the hill peoples of Assam. Ann. Missouri Bot. Gard. 36: 355-404, Sut6, T. and Y. Yoshida, 1956. Characteristics of the Oriental maize, Jn Land and Crops of Nepal Himalaya II: 373-529. Vavilov, N. I., 1931. Mexico and Central America as the prin- cipal center of origin of cultivated plants of the New World. Bull. Appl. Bot. Gen. Pl. Breeding 26: 179-199. Weatherwax, P., 1923. The story of the maize plant. Univ. Chicago Press. ——-, 1936. The origin of the maize plant and maize agriculture in ancient America. Univ. New Mexico Bull. (October), 1-8. ——-, 1954. Indian corn in old America. New York, Macmillan. Wellhausen, E. J., A. Fuentes O. and A. Hernandez C., in col- laboration with P. C. Mangelsdorf, 1957. Races of maize in Cen- tral America. Nat. Acad. Sci.-Nat. Res. Council Publ. No. 511. Willey, G. R., 1955. The prehistoric civilizations of nuclear America. Amer. Anthrop. 57: 571-593. Wright, A.C. A., 1949. Maize names as indicators of economic contacts. Uganda Jour. 13: 61-81. THE ORIGIN OF CORN V. A Critique or CuRRENT THEORIES BY Rosert G. REEVES AND Paut C. MANGELSDORF In the previous papers in this series (22, 23, 24, 32), we have reviewed the objections to and the evidence sup- porting our tripartite theory on the origin and evolution of corn. We have shown that the factual evidence for some parts of this theory has increased substantially dur- ing the last two decades, and that nothing has yet come to our attention which completely rules out any part of it. The hypothesis that the original corn was a type of pod corn is all but proven by archaeological evidence and by a genetic reconstruction of the ancestral form (15, 16, 17, 19, 20). Because of the discovery of fossil corn pollen in Mexico, the view that corn originated in South Amer- ica is less satisfactory now than it was twenty years ago, but the hypothesis of a South American origin is not yet completely untenable since research on some phases of this problem, such as that on fossil pollen, has only begun. The evidence that teosinte (Zea mexicana) originated as a hybrid between corn and ‘Tripsacum is decidedly stronger now than when the hypothesis was first pre- sented and, contrary to opinions of several other workers, there is no sound evidence against the hypothesis. We have always recognized that this part of the tripartite theory cannot easily be tested. We have never consid- [ 428 ] ered the probability to be great that a ‘‘good’’ teosinte could be synthesized by hybridizing corn and Tripsacum in experimental cultures, and we have given reasons for this. The recent evidence is decidedly in favor of the view that introgression between corn and teosinte has been frequent and that it has been effective in producing in- numerable new varieties and forms, both ancient and modern. We now consider that this third part of the tri- partite theory is almost an established fact. It should be clear from the contents of previous papers in this series that we consider the tripartite theory to be better supported by factual evidence than any other ex- planation of the origin of corn proposed up to the present. The fact that we are still committed to the tripartite theory, however, should not prevent us from consider- ing alternative theories and this we shall attempt to do objectively, if briefly, in this final paper. THE PApyRrescCENT (Semivestidos) THEORY This is no more than a slight modification of the pod- corn theory. Andres (1) discovered in Argentine maize a type which superficially resembles a weak form of pod corn. Apparently unaware that Bonvicini (5) in Italy had described this character many years earlier and had given it the name ‘‘palee sviluppate,’’ Andres called the type ‘‘semivestidos’’ and suggested that it, rather than pod corn, might be the ancestral form. The character has recently been given still a third and probably more appropriate name ‘‘papyrescent’’ by Gal- inat (10), whose studies show that the glumes of this type become soft and papery as they mature. Unlike pod corn, which although sometimes monstrous still represents a combination of normal characteristics found in other grasses, papyrescent is a defect in development which it [ 429 ] is difficult to regard as constituting the primitive form of modern corn. No archaeological specimens of papy- rescent corn have been reported. As was pointed out in the first paper in this series, Weatherwax (41), in discussing one aspect of the pod- corn theory, apparently confused papyrescent maize and a weak form of pod corn, half-tunicate. His illustration (Fig. 51) of half-tunicate maize is almost certainly a photograph of papyrescent maize. THe Corn Grass THEORY Singleton (35) has suggested that the ancestral form of modern corn is ‘‘corn grass.”’ This anomalous type, the product of a single dominant gene, produces numer- ous tillers and small ‘‘ears’’ with a high proportion of single spikelets. Many of the kernels are partly enclosed in bracts, but the majority of these are not glumes but spathes. He also suggested that, if a plant of corn grass were found in nature, it would not be recognized as maize and would almost certainly be regarded as a different species if not a different genus. This may be true, and it illus- trates how the maize plant can be drastically changed by a single gene mutation. If corn grass were the ancestral form, a mutation at a single locus could have transformed it from a wild, almost useless, plant to the unique cereal which maize is today. Although corn grass has some of the characteristics which we might expect to find in an ancestral form — for example, a freely-tillering habit —it lacks others, such as the regular development of prominent glumes. At the other extreme, it has characters which are not demanded of a hypothetical ancestor. One of these, single spikelets (9), represents a condition more specialized in- stead of more primitive than the paired spikelets of mod- [ 430 | ern maize. Another, a well-developed spathe, suggests the ancestral form not of maize but of Cow, whose fruit case has been found by Weatherwax (39) to comprise a spathe and a segment of the rachis. Corn grass probably is, as Galinat (8) has suggested, a ‘‘false’’ progenitor of maize, exhibiting certain traits which might have oc- curred in a remote ancestor of the Maydeae. Finally, the evidence from archaeological maize does not support the corn grass theory. Prehistoric maize had prominent glumes, but it did not have the long spathes of corn grass. The possibility that corn grass is the an- cestral form appears to us to be remote indeed. THE TEosinvTe THEORY The theory that maize originated as a domesticated form of teosinte—its nearest known relative— was first proposed by Ascherson (2). Later students, notably Harshberger (12) and Collins (6), modified the theory postulating that one parent of corn is teosinte and the other isa grass now unknown. As teosinte occurs natu- rally only in Mexico and Central America, supporters of this theory have usually assumed that both teosinte and maize originated in that region. We (21) concluded that teosinte is the progeny rather than the progenitor of maize — the product of the natural hybridization of maize and its wild relative, Tripsacum. ‘Teosinte, how- ever, plays an important role in the tripartite theory, for this theory holds that the many modern varieties of maize are the product of the introgression of teosinte into maize. Since 1989, new evidence has been presented both in support of and in contradiction to the teosinte theory. Beadle’s (4) discovery that the seeds of teosinte will *‘pop’’ when exposed to heat, shattering the hard, bony shell in which they are enclosed, shows one way in which teosinte might have been used as a food plant and weak- [ 431 ] ens the objection that a species so unpromising for food purposes would never have been domesticated. There is no evidence, however, archaeological, historical or con- temporary, to show that teosinte was ever employed for food in this manner. When teosinte is used for food, as it occasionally is today in times of food shortage, the fruits are crushed ona metate or with a mortar and pes- tle, and the meal of the crushed caryopses is separated from the fragments of the bony fruit case.’ Langham’s (13) data on the inheritance of character- istics which distinguish teosinte and maize indicate simple Mendelian inheritance for several characters and lend some support to Emerson’s (unpublished) contention that a few large scale mutations could transform teosinte into maize. But the much more extensive data of Man- gelsdorf (14) and of Rogers (38, 84) show that the genes which distinguish maize and teosinte are numerous and are distributed among a majority of the chromosomes. The highly significant studies of Rogers seem to have been completley overlooked by both Weatherwax and Randolph; at least they are not cited in their extensive bibliographies. The recent studies of Barghoorn, Wolfe and Clisby (3) on fossil pollen in Mexico lend no support to the teosinte theory. Although pollen of both maize and Tripsacum was found at great depths, the pollen of teo- sinte occurred only near the surface in the upper levels of the drill core where maize pollen was abundant sug- gesting that the practice of agriculture had begun. Furthermore, the maize pollen found at the lower levels is as large as any modern maize pollen and shows no re- semblance to teosinte pollen in the ratio of total diameter to the diameter of its pore. If this fossil pollen is as old as it is estimated to be — 80,000 years or more — the ' Personal communication from the late R. H. Barlow. [ 432 ] theory that maize originated from teosinte under domes- tication can now be safely ruled out. Recent studies of archaeological maize, like those of fossil pollen, do not support the teosinte theory. On the contrary, they show that the earliest maize was less like teosinte, whereas some recent maize is more like it. Archaeological specimens exhibiting characteristics of teosinte, including distichous ears, single spikelets and highly lignified rachises and glumes, have been found in several sites. But these are always recent specimens and are interpreted as being products of the introgression of teosinte into cultivated maize (7, 11, 19, 20, 25). A series of studies on the morphology of the corn ear has a bearing on the teosinte theory, because many work- ers who favored this theory explained the polystichous character of the corn ear as the result of the lateral fusion of several teosinte spikes. The voluminous literature on this subject was reviewed by Mangelsdorf and Reeves in 1939 (21) and more recently by Nickerson (27). The present status of the problem is that evidence for the lateral fusion of two-rowed spikes to form the polysti- chous ear is completely lacking; the only evidence found for fusion is the adnation of the rachis flaps (prophylls) to the main axis of the cob. It may be concluded, there- fore, that the structure of the corn ear has thus far shown no evidence that corn is a descendant of teosinte. THe ‘THeory or COMMON ANCESTRY It appears that Montgomery (26) was the first to pro- pose the theory of common ancestry, although he did not include Tripsacum in the alliance with corn and teo- sinte. Weatherwax (87) formulated the theory as we now know it, by adding Tripsacum to the two species considered by Montgomery, and he defended it in sub- sequent publications (88, 40, 41, 42). Randolph (28, 29) [ 433 ] agreed with Weatherwax, with reservations; he still re- gards the direct descent of corn from teosinte as a dis- tinct possibility. The theory of common ancestry maintains that corn originated from a perennial, wild, corn-like ancestor, now extinct, and that this extinct ancestor, sometimes called pre-maize, in turn had an ancestor, likewise extinct, in common with teosinte and Tripsacum; also that the na- tive range of all of these species was Central America and Mexico. Actually the theory represents the application to the American Maydeae of the broad views of Darwin and earlier students of evolution. The factual evidence claimed by the proponents of this theory falls into two categories. (A) The three groups— corn, teosinte and Tripsacum—are very similar, except that each has modifications of its own which have led to the differences now found among them. For example, teosinte and Tripsacum have lost one member of each original pair of pistillate spikelets, but corn has not; most varieties of corn and teosinte have lost the terminal stam- inate portion of the lateral inflorescence (ear), but Trip- sacum has not. Weatherwax (41, 42) pointed out by way of explanation that, if we could restore to each of the three groups the primitive organs which have been mod- ified in evolution, they would converge in a common type, giving us an idea of the common ancestor. But more revealing, in our opinion, is the result that would be obtained by restoring the primitive organs of only corn and Tripsacum, omitting teosinte. ‘The common type towards which they would converge is the same as when teosinte is included. This fact is best explained by the hypothesis of the hybrid origin of teosinte, because this hypothesis holds that the characters of teosinte are merely a combination of those of corn and ‘Tripsacum. (B) It is stated that corn, teosinte and 'Tripsacum are [ 434 ] now sympatric only in Central America and Mexico, and for this reason all of them probably originated from a common ancestor which also occurred there. When the fallacy of placing such strong reliance on this present-day distribution is recognized, little or no factual evidence for the theory of common ancestry remains. Stebbins (36) reviews the literature on the theory that centers of diversity correspond to centers of origin. He points out that the theory has many pitfalls, except when the group in question is young and the selective forces of the en- vironment have been operating in about the same man- ner throughout its evolutionary history. Stebbins shows that related genera, congeneric species and even conspe- cific populations might be expected to have widely dis- junct ranges. He cites numerous examples of a single species with two ranges separated by half the distance around the earth. He shows also that examples of closely related disjunct taxa are not rare exceptions but are nu- merous and that some of them have been known since the time of Darwin. Much of the validity of the Montgomery-W eatherwax theory depends upon the assumed correlation between the common ancestry and the common place of origin of the three groups. If one or two of the groups were shown to have originated elsewhere than in the present center of diversity (and this may yet prove to be true especially of Tripsacum), this would seriously weaken the theory. In addition, the proponents of the theory assume the previous existence of a pre-maize and of the common ancestor of pre-maize, teosinte and Tripsacum without one iota of evidence. The tripartite theory, in contrast, is more flexible and much less dependent upon com- pletely unknown ancestors of corn. Indeed, it requires no ancestral types other than forms still in existence. Types of pod corn very similar to our hypothetical an- [ 435 ] cestor occur today; types of Tripsacum and corn which come extremely near to satisfying the requirements for the putative parents of teosinte are well known. Yet Weatherwax (41,42) states that it is the tripartite theory, rather than the “‘simple’’ theory of common ancestry, which ‘‘is topheavy with assumptions of such character that if one of them should be rejected the whole struc- ture would fall.”’ The theory of common ancestry has two additional weaknesses which are serious: (A) It does not, Weather- wax’s and Randolph’s contentions to the contrary, not- withstanding, explain all of the known facts. (B) It can not easily be tested. A. Some of the facts which the theory of common ancestry does not explain are discussed in detail in other papers of this series (28, 32). Here it will suffice to point out that the theory does not account for the facts that (a) teosinte is intermediate between maize and Tripsacum in a great majority of its characteristics (21, 31); (b) early archaeological maize is more ‘‘maize-like’’ than later maize (7, 11, 19, 20, 25); (c) fossil pollen of maize and Tripsacum were found at great depths at one site in Mexico, whereas teosinte pollen occurred only in the upper levels of the drill core; (d) forms of pod corn are now in existence which possess all of the characteristics expected in the ancestral form (16, 17); (e) variation in knob numbers is correlated with tripsacoid characteristics (18), and with proximity to Guatemala, the reputed center of origin of teosinte (21, 80). B. The theory is largely untestable, because the only evidence which could prove it to be correct beyond a reasonable doubt would be the discovery of prehistoric remains, antedating agriculture, of all three groups of the American Maydeae and of the remote ancestor from which these three groups stem. Since it is largely untest- [ 436 ] able, it neither stimulates new research nor points to possible new methods of maize improvement. In this respect, the theory is less useful than the tripartite theory which has furnished the impetus for an extensive series of researches on maize and its relatives and has also sug- gested new possibilities for improving maize. If two theories appear to be equal in validity, the one which is testable and which stimulates new research is the more useful; a theory which is plausible but untestable tends to stifle research. In emphasizing the differences between the tripartite theory and that of common ancestry it should not be overlooked that there are also important resemblances between them. The two theories agree that (a) corn is an American plant; (b) it is descended, with Tripsacum, from a remote common ancestor; (c) its immediate an- cestor was a freely-branching plant bearing small ears with grains enclosed in glumes; (d) corn attained its present form through changes occurring during domes- tication, which began not more than a few thousand years ago. Ina broad sense, then, the two theories agree with respect to the place, time and manner of origin. In the same broad sense, the problem of the origin of corn can almost be said to be solved. [ 437 ] to ~ 10, ll. 12, 13. LITERATURE CITED . Andres, J. M., 1950. Granos semivestidos, restos de un caracter ancestral del maiz. Revista Argentina de Agron. 17: 252-256, Ascherson, P., 1875. Ueber Fuchlaena mexicana Schrad. Bot. Vereins Prov. 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