BOTANICAL MUSEUM
LEAFLETS
HARVARD UNIVERSITY

PRINTED AND PUBLISHED AT THE
BOTANICAL MUSEUM
CAMBRIDGE, MASSACHUSETTS

BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY

VOLUME XXVI

BOTANICAL MUSEUM
CAMBRIDGE, MASSACHUSETTS
1978

DATES OF PUBLICATION - VOLUME 26

INGOs, 0 see oe fo cttceee caren Sec eancvreuses ues 19 ceg bites dara oeesscatmsiay4 October 19, 1978
INOSs 25 Arie ecsanion dese teatedienstes seen guetisiteasg siameerrndenad November 16, 1978
INI 29s ceseee sos trate ce ee been pag etaehmecsaldatdeaceosslantecnae saints December 20, 1978
INGOs Gree oe pcdsiatresscteseks pisealsees ee ueice er aceueupenigecssesencse February 8, 1979
INO 5 Fete teeter paca start oon eeaedetseah aseuing eset anen penn March 13, 1979
NOs 8 wien ececesdes oiwwccewnncrwesdaendensseved seaneipandéaeiaberwinesadas May 20, 1979

DNOSs. 902: TO" ces ens sees eeccaccavicecgscacoueeecineune ois oierenasamech November 29,

1979

TABLE OF CONTENTS

NUMBER | (Jan. 31, 1978)

Studies in American Orchids X
DW IESE A CIABAY cc sertivenideaeeanitenpsanuvecsaisvscess l

Revision of the Genus Masdevallia - |
by HERMAN R. SWEET. ...........cccccceececceceeccececeeaeens 39

Studies in the genus Oncidium - II. Section Serpentia
Oy OPIN, Foe ERO ies cee oeinccy eastern reaececues nausea cceess 55

NUMBERS 2 - 4 (Feb.-April 1978)

Bibliography of Precambrian Paleontology
and Paleobiology
by STANLEY M. AWRAMIK AND
EO PUN RGOORS eeu viviseensod ceieeednceses elas eenaaias 65

NUMBER 5 (May 30, 1978)

De Plantis Toxicariis e Mundo Novo Tropicale Commen-
tationes XXIII. Notes on Biodynamic Plants of
Aboriginal Use in the Northwestern Amazonia
by RICHARD EVANS SCHULTES ......cc.cceeceeeeeeeeeeeeeees ye

Cocaine in Blood of Coca Chewers
by B. HOLMSTEDT, J.-E. LINDGREN, L. RIVIER AND
Be IAN acoso thse g senna ran trsesteaataueeeteiserai os 199

Chromosome Numbers in Neotropical Erythroxylum
(Erythroxylaceae)
by T. PLOWMAN, L. RUDENBERG
PI Ss WIRE EG du dirrenciutaeuusarersaatenierseebarniccediawe: 203

NUMBER 6 (June 30, 1978)

Soma Brought Up-to-date
by R. GORDON WASSON .........ccc cece ceceeceesceeseeeeeeeees 211

De Plantis Toxicariis e Mundo Novo Tropicale
Commentationes XXIII
DY RICHARD EVANS SCHULTES: ccccscessekicsiscaivaviveuace 225

NUMBER 7 (Sept. 30, 1978)

Fossil Pollen and the Origin of Corn. Ancient Pollen from
Deep Cores in Mexico Shows the Ancestor of Corn to be
Corn and not its Relative, Teosinte
by PAUL C. MANGELSDORF, ELSO S. BARGHOORN
AND UMESH C. BANERJEE ....cc.scccceereccessensstoseenacens 234

NUMBER 8&8 (Oct. 31, 1978)

Effects of Field Preservation on the Flavonoid Content of
Jessenia Bataua
by GILLIAN A. COOPER-DRIVER AND
MICHAEL. al. BAGICR. -csssadsescevicecsass aanchews test parerar es 257

NUMBERS 9 -10 (Nov.-Dec. 1978)

A Neglected Mayan Galactagogue. Ixbut

(Euphorbia lancifolia)

by FREDERIC ROSENGARTEN, JR. ......scsscecseeeeesneeeeees 277
Evolution of the Identification of the Major South

American Narcotic Plants

by RICHARD EVANS SCHULTES: sccsasecisnssarnoreermeateees 311
The Correct Name for Siberian Ginseng

by DyaAJA D. SOEJARTO AND

NORMAN R. FARNSWORTH ceccacsatiiees etactseerterresremeee JOP

V1

INDEX OF ILLUSTRATIONS

PLATE
BTA CI GE PONCE. eit occ aniaeies tak bs ucass eaten seaeuawieadeess: 4
Chonidrorhyncha Stenioides ....ccccccccccccccccceccccucccuseccuceens 7
Cybedbus QrANAIS ...ccccc cece ccccccccccccccccccccececccecccceccescaceees 3
Cyclopocon OVGUPONUM ciiccccicasevscicnscssssssesotsisecasdeceseess 3
PE CARINES FOVINOSUS céiccsiaiinesxcurinnceiinesenieariernenidarvateds 2
FUCHS RNTDIC cevcslavessescscsncesseevasexiadeanccacduessadaseaese: ps
Eliroplectris PQUCULOLG vicisiciccacacsesevs vndescvncesvssssecvansensess 5
Epidendritnt trOpidiOideS is cciivsivacsascinscavssacsenssavecersrereres 6
Erythroxylon Coca and novogranatense,

microsporocytes and root tip cells of ...................000es 17
Euphorbia lancifolia (xbut) ........cc cece cece cee eceeeeeeeeueees 29

EPOWIR IE TX PUL LOW. <censceensraxcchdacscucteniubusdadnasieanaxvncds 34

POTS! TET 4 x covesriusewinseastcsiakas chase erasels 24 ,27,30,31

Be rac ede eo ea rc eee dieneenneens 25

Leaves showing preparation of tea ..................0.08. CAB

Map of Guatemala and adjacent countries

SHOWING FANGe OF IXOUL, csi sciides ca vidiasenuesvecicdseoenee 28

oN 111) ce oe ee eae eR ra nats wey Tey nen eee eee eee rere 26

Young mother nursing baby after having

CARON POU CED. yc tencreercrcisanavcisaniacacent seein)
FIGRGIGKCHIS LINGIOVQMUS iasse ccvcuscdvidetcnainscddaaniaiest aes 4
Jessenia Bataua, near Manaus, Brazil .................6..0.0 66 20
PUSTLCIG DOCTOVOIIS: oc.0ciewckxca cd rervexendbisvevianatensdeinbevaeexs 21
Justicia pectoralis var. stenOphylla ciccccccccccccccccccccceeeees 22
Kunth’s report of the source of the snuff,

UR, 2 cere tsa eres eda acisedcatey sens ee eee 38
MASAOVAING BOMOU xdcsiecticedascancrkritakehVidedsedstnsesiveieters 10
MGSACVAING CHOME secstccerctpitrnucasntsisewsnsetinsetiveesnannt 8,10
MGASAEVQIIAIADGNG: jrceisararacivecersxisuteersseraisisiansixesauors 9
Masdevallia kypNonantha .....cccc cece cc ccc cen nc ccc enneeenneees 8,12
Masdevaliia: LARSDOCVOU. cic cecaxcvesavatesecatanteexicsabsestotases 1]
Masdevallia pSeudOMinuta ......cccccc cece cece nc eneeeneseeneees 8,12
WEAN OVOIE EO TIIOON: occa ccsecadev iia tts cect reguaes eases 10
Masdevallia veneZzueland .......cccceccccceccecsccccccccccesseceess 13
NECHAOHOTA GSD cic wcankitatcucriwedveunktsceneiwaxaeeswiesewedion 23
OnCiGIUIN BIGDUN PONE: oi sisaxcncicedaninucsdeencisecccvavncisienees 16
CPHCUGIONR TIOPHGE, osc ccc asta cacevdcietuns axvssinweeeatinedencewes 15.16

Vii

ORCTUIHIA SCAANOP cesesaceconecasausestie tikcleaveceas irene 14

ORCA SOT OPUS: escassridaceseeeesesdaceerey nee eer eee 16
OSIMOGIOSSUl DANGHIGIUDE ssidcsvssuass en teteevnsdasestena tin etaiawes 7
POWHOFCH SCOIOTUIAONG. ~ncioatciae in dindesesvieiasiaresectueyensseis 5
OIG FIT pcs wc ce icone tnd eax casawitesnnabeareca tease 4
PAP INCIVS. POPUVIONY. ciicestaicerssuececesdin se eecsds daseeniasarnes 5

Pollen grains of Mexican popcorn, and
archeological pollen grains ca. 1600, and ca.

PO AGS OIE | oecnsaeosdovencencuueiensecroienerianeeasaaedtaies 18
PSCUAGCENINUIN PUIG cco sshccsuuvescxdexsinacaay eda denenies tatieies l
PS COS, 6 ince cies ue cnyadens cus wihene so pohrwenind |
Report of jurema by Morais in 1881] .......... 0. eee eee eee eee 39
Report of Philippi of toxic nature of Latud ........ ccc 37
ROOST STR: cesta ineasnats neers sess ecdseeenteeer ees 3
DOPCOCIOINS DENQIOIGUE cessenientstiveussdicyenqenitiaseciteab vere 4
Villavicencio’s description of the use of

G@vandascd Wi PCUadOr CIS58)) ccocientcceciariirieeuieiteast 4()

Von Humboldt’s identification of the source
of the narcotic snuff, niopo,
of the Otomac INdians. ..cssdncciwosivesnavcsavndessesecawernaaee’s 36

Vill

Acanthaceous plants
Northwest Amazon
ACANTHOPANAX
eleutherococcus
senticosus 339
ACOSMIUM
nitens 230
Aja Ekapad 220
ANADENANTHERA
colubrina 319
peregrina 319
a-nama-he-se-he-pa
ANAXAGOREA sp.
ANTHURIUM
crassinervium
var. caatingae
APHELANDRA
aurantiaca 267
pilosa 268
arbol de los brujos 328
atiipa-kashaikie-si-sehe-pa
ayahuasca 192,322,337

BANARA
guianensis 234
BANISTERIOPSIS
Cabrerana 327
Caapi 327
inebrians

Martiniana
quitensis
Rusbyana
bawn’-tez
beetle tree
BOMBAX
globosum
bov-fee’ 197
BROSIMUM
utile 180

BUCHTIENIA
rosea 23
BLETIA

capitate

267

342

ba

wet
228

179

399

327
327
327
See
181
pt

937

2

caapi 322
CARAIPA
parvielliptica

tr

180

INDEX

CHONDRORHYNCHA
guttata 27
stenioides 26
chromosome numbers
Erythroxylon Coca 206,207
Erythroxylon emarginatum
Erythroxylon Kunthianum
Erythroxylon Mannii 206
Erythroxylon novograna-
tense 207
Erythroxylon orinocense
chuftanguemesha 236
chu-ru-ko-pu 270
cocaine in blood
cohoba 317
COMBRETUM
Cacoucia 235
conoco 186
corn
fossil pollen of 239
two-eared husk system
COSTUS
aMazonicus
erythrocoryne
COUSSAPOA
cinnamomea
magnifolia
var. glabrescens
orthoneura 181
trinervia 182
CROTON
glabellus 232
palanostigma
cupuassu 186
CURAREA
tecunarum
CYBEBUS 15
grandis 15
CY CLOPOGON
lineatus 20
ovalifolium 16,17
stenoglossus 20

CYPHOMANDRA

206
206

207

199

244

179
179

181

181

232

228

16

crassifolia 191
dolichorachis 19]
endopogon 191

DICHAEA
muricata 180
do-ro-he 235

ELEUTHEROCOCCUS
koreanus 342
senticosus 339
ELLEANTHUS 6
brasiliensis 8

var. Hookeranus 10
capitatus 8,9

Caroli 12
casapensis 9
cephalophorus 8
cephalotus 9
confusus 13
cynarocephalus 9
formosus 14
Glomera 10
Hookeranus 10
Killipti 11
lancifolius 6
lateralis 12

Sect. Calelyna 12
Sect. Cephalelyna 7
Sect. Chloidelyna_ 13
Sect. Elleanthus 14
Sect. Hymenophora = 12
Sect. Laterales 12
Sect. Otiophora 12
Sect. Stachydelyna 13
Sect. Virgatae 14
Sodiroi 11
sphaerocephalus 12
Subg. Calelyna 12
Subsect. Chloidelyna 13
ELTROPLECTRIS
pauciflora 24
epena 270
EPIDENDRUM
capttatum 9
trigoniflorum 25
tropidiovides 26
ERYTHRODES
pauciflora 24
ERYTHROXYLON 312
Coca 199,203
EUCHARIS

amazonica 178
EUPHORBIA

lancifolia = 277

Evelyna

brasiliensis 8
capitata 8
casapensis 7,9
cephalophora 8
cynarocephala 9
Hookerana 10
hymenophora 12
myrosmatis 12

Sect. Cephalelyna 7
Sect. Stachydelyna 13
Subg. Calelyna 12
Subsect. Chloidelyna = 13

FITTONIA
argyvroneura 268
Verschaffeltti 268

Galac-latex 285
galactagogue 277
Ginseng, Siberian 339
GLOMERA
brasiliensis 10
GUATTERIA
Duckeana 229
dura 229
GYROSTACHYS
lineata 20
ovalifolia 17

HABENARIA
Ernestil |
linguicruris
HAPALORCHIS
Lindlevanus 21
lineatus 20
tenuis 20
han-shee-ma_ 182
HEDERA
senticosa 342
HEDYCHIUM
coronarium — 180
HELICOSTYLIS
scabra 182
HERRANIA
Camargoana 233
HETEROSTEMON
mimosoides 231
HEVEA
brasiliensis 185
guianensis 185

INSERIA

rosea 195
IRYANTHERA
macrophylla 316
ixbut 277

JESSENIA
Batawa 257
JUANULLOA
ochracea 191
jurema 336
JUSTICIA
Blackei 268
Cabrerae 269
chlorostachya 269
comata 269
ideogenes 269
pectoralis 270
var. stenophylla 270
Schultesii 271

ka-ne-wee-re 233
ka-sa-ra 235
ka-we’-re-ta 188
ké-re’-ma_ 269
kon-si-ahipa-cho 179

LATUA 321
pubiflora 328
latue 328
LYCOPODIUM

cernuum = 225

MABEA

nitida 232
ma-ha-shu-ke-ma_ 195
ma-ree-m'a_ 181

MARKEA
coccinea 192
MASCAGNIA

glandulifera 327
MASDEVALLIA
Bangti 40,41

gnoma 41

irapana 42
kyphonantha 44
Lansbergti 43
pseudominuta 45
Sect. Triotosiphon 40
Subg. Triotosiphon 40
trioon 46
venezuelana 47

XI

mata-cachorro 183
ma-tsé’-ko-ra_ 180
MAXILLARIA
pantherina Rchb.f. 28
pantherina Hoffmsegg. 28
pardalina 28
MAYNA

longifolia 234
toxica 235
mee-kee-na-ta-ree 269
mee-na-ku-ru 268
mee-o 182
mee-re-ka-sé-pa 270
MENDONCIA
aspera 271
MICRANDRA
Spruceana 185
MILTONIA

anceps 30

flava 30

MIMOSA 319
hostilis 322
verrucosa 322
minakoro-se-he-pa 268
mun 182

MYRICA

parvifolia 227

natema 322
NIPHOGETON
ternatus 188
non-ne 221
nyakwana 270

OCTOMERIA

truncata 25
ODONTOGLOSSUM
anceps 30
angustatum

var. heterosepalum 31
candidum 28
heterosepalum 31
ONCIDIUM

anceps 30
convolvulaceum — 57
globuliferum 57

var. costaricense 57
Harlingti 58
pusillum 180
Scansor 59

Sect. Serpentia 56
serpens 56,59

trachycaulon 59
Wercklei 57
00-too-mee-ko 235
OSMOGLOSSUM
candidum 28
panduratum 29
OSTEOPHLOEM
platyspermum 230
OXALIS

lotoides 231

PAGAMEA

coriacea 195
macrophylla 195
paleobiology, bibliography of
Precambium 65-175
paleontology, bibliography of
Precambium 65-175
PALMORCHIS
colombiana 2
PELEXIA

corymbosa 19
Hameri 22
pauciflora 24
PEPEROMIA
macrostachya

var. nematostachya 226
PHYSOSIPHON
Bangi 41
Lansbergti 43
PHYSURUS
pauciflorus 24
pinde 322
PIPER
hostmannianum 226
interitum 226
PLATYTHELYS
peruviana 24
POGONIA
physurifolia 3
Raventi 3
po-ham-bo’-raek 183
popcorn 243
PORPHYROSTACHYS
parviflora |
POTALIA

Amara 189
POTHOMORPHE
umbellata 227
POULSENIA

armata 182

XII

POUROUMA

Schultesii 182
Precambium paleobiology,
bibliography of 65-175
Precambium paleontology.
bibliography of 65,175
PSEUDOCALYMMA
alliaceum 236
PSEUDOCENTRUM
Purdii 4
PSEUDOLMEDIA
laevigata 182
PSILOCHILUS
carinatus 2
physurifolius 3
PSYGMORCHIS

pusilla 180

putika 217

putka 217

QUALEA
acuminata 231]

RETINIPHYLLUM
concolor 196
pilosum 196
Schomburgkit 196
speciosum 196
truncatum = 197

ROUREA
cuspidata 183
glabra 183

RUELLIA
colorata 271
RUMEX
Obtusifolius 227
RYANIA

angustifolia 235

SANCHEZIA
thinophila 272
sapillo 280

SARACHA
procumbens 236
SARCOGLOTTIS

Lehmannii 22
SAUROGLOSSUM
andinum 18
aurantiacum 18
corymbosum 19
distans 19

monophyllum 20
tenue 20

SCHIEKIA
orinocensis 178
SCHOENOBIBLUS

peruvianus 187
see-tu-ku-"he-ta 271
se-ne’-no 183
shahasi-sehé’-pa_ 180
snuff 315
SOBRALIA
Amparoae 5
Fenzliana 4
Hawkesti 5
luteola 5
panamensis 4
pleiantha §
Powellii 5
Warscewiczli 5
SOLANUM
albidum 192
apaporanum — 192
campaniforme 193
jJamaicense 193
mammosum 193
subinerme 194
Topiro 194
verbascifolium 194
soma 211

color of 219
etymology of 219
reviews of 223
sorcerers’ tree 328
SPIRANTHES
amabilis 20
Fawcettii 20
lineata 20

nitida

var. andina 18
Preslii 17
subumbellata 19
tenuis 20
Weberbaueri

var. aurantiaca 18
STELIS
ophioglossoides 25
trigoniflora 25

STENOPTERA
parviflora |
pilifera

var. parviflora |

XH

STYRAX
Tessmannit 188
yvapobodensis 188
SYMPHONIA
globulifera 234
SYNASSA
corymbosa 19
dilatata 19

TACHIGALIA
cavipes 183
paniculata

var. comosa 185
ptychophysca 185
TELIOSTACHYA
lanceolata

var. crispa 272
TELITOXICUM
peruvianum 228
teosinte 237
TETRAPTERIS
methystica 328
mucronata 328
tetsi 227
THEOBROMA
grandiflorum 186
subincanum 187
TRIPHORA
Raventi 3
TRIPSACUM 242

UNONOPSIS

veneficiorum 229

vinho de jurema 319

VIROLA 313
albidiflora 183
calophylla 316
calophylloidea 316
cuspidata 316
elongata 316
loretensis 316
Pavonis 316
rufula 316
surinamensis 316
theiodora 315,316

VISMIA

ferruginea 234

wa-so-gee_ 181

XYLOPIA ya-poo-moo-ho 235
amazonica 229 yaje 322
Benthamii 230

XIV

BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY

CAMBRIDGE, MASSACHUSETTS. JANUARY a; 1978 Vor. 26, No.

STUDIES IN AMERICAN ORCHIDS X*

LESLIE A. GARAY

In preparing the final manuscript of the Orchid Flora of
Colombia and Ecuador a number of correlated material from
the neighboring countries had to be investigated. These studies
which cannot directly be incorporated into the main text of the
‘*Flora’’, are presented here. Number IX of this series was
published in Botanical Museum Leaflets, 24: 299-304, 1973.

Habenaria linguicruris Rchb.f., Bonpl. 2: 10, 1854.
Type: Venezuela; Caracas, WAGENER 96! (W).
Syn.: Habenaria Ernstii Schitr., Fedde, Rep. Beih. 6: 26,
1919.
Type: Venezuela; Caracas, ERNST s.n.! (AMES).

A comparison of the holotypes of H. linguicruris and H.
Ernstii with one another has shown them to be conspecific.

Porphyrostachys parviflora (C. Schweinf.) Garay, stat. nov.
Basionym: Stenoptera pilifera var. parviflora C. Schweinf.
in Bot. Mus. Leafl. Harv. Univ. 9: 222, 1941.
Syn.: Stenoptera parviflora (C. Schweinf.) C. Schweinf. in
Fieldiana Bot. 33: 4, 1970.
Type: Peru; Huanuco, Llata, MACBRIDE & FEATHERSTONE
2301! (F).

*This study is supported by a grant DEB 73-01354-A03 of the National Science
Foundation which is gratefully acknowledged.

Published monthly except during July and August by the Botanical Museum, Harvard University,
Cambridge, Massachusetts 02138. Printed by Harvard University Printing Office. Subscription:
$20.00 a year, net, postpaid. Orders should be directed to Secretary of Publications at the above
address. Second-Class Postage Paid at Boston, Mass.

7

The plants of the genus Porphyrostachys may be separated
from Stenoptera in that the flowers have a column with a long.
decurrent foot to which the lip is adnate; lateral sepals are also
decurrent on the ovary. These characters are absent in Stenop-
lerd.

Psilochilus carinatus Garay, sp. nov.

Terrestris, erecta, usque ad 30 cm. alta; caulibus ascenden-
tibus, pluriarticulatis, primum vaginatis, deinde laxe paucifo-
liatis; foliis ovato-ellipticis, acutis vel subacuminatis, basi
rotundatis, breviter petiolatis, usque ad 9.5 cm. longis, 4 cm.
latis; inflorescentia terminali, pauciflora; bracteis distichis,
ovato-lanceolatis, usque ad 1.5 cm. longis, sursum decrescen-
tibus; floribus succedaneis: sepalo postico lineari-oblongo,
acuto, dorsaliter in medio carinato-incrassato, usque ad 21
mm. longo, 3.5 mm. lato; sepalis lateralibus falcato-linearibus,
acutis, dorsaliter subcarinatis, usque ad 18 mm. longis, 3.5
mm. latis; petalis tenuibus, falcato-oblanceolatis, acutis, usque
ad 18 mm. longis, 3 mm. latis; labello longe unguiculato, deinde
cuneato-trilobo, lobis lateralibus oblique triangularibus, lobo
intermedio antico, subquadrato-rotundato, antice truncato,
margine eroso-denticulato, disco in medio bicarinato, Carina
tertia in ungue, usque ad 17 mm. longo, 7 mm. lato; columna
gracill, arcuata, alis subtruncato-denticulatis, usque ad 15 mm.
longa; ovario cylindrico, breviter pedicellato, usque ad 15 mm.
longo.

Colombia: Sierra Nevada de Santa Marta, PURDIE s.n.!
type! (K); Fusagasuga, HOLTON s.n.! (K).

Palmorchis colombiana Garay, sp. nov.

Herba terrestris, elata, pars quae adest, 40 cm. longa; radici-
bus nondum notis; caulibus supra foliosis; foliis satis tenuibus,
ellipticis, acutis vel abrupte acuminatis, basi longe petiolatis;
lamina usque ad 20 cm. longa, 7.5 cm. lata; petiolo usque ad 10
cm. longo; inflorescentiis satis gracilibus, remote plurivagina-
tis, supra subdense plurifloris, usque ad 12 cm. longis:; bracteis
lanceolatis, acuminatis, usque ad 10 mm. longis; floribus al-
bidis, succedaneis; sepalo postico lineari-ligulato, obtusius-

9

-

culo, usque ad 12 mm. longo, 2 mm. lato; sepalis lateralibus
oblique falcato-oblanceolatis, obtusis, usque ad 10 mm. longis,
2.5 mm. latis; petalis oblanceolato-dolabriformibus, obtusis,
usque ad 11.5 mm. longis, 3 mm. latis; labello basin lineari-
conduplicato, puberulo, deinde trapezoideo, utrinque ob-
tusangulo, antice tridentato, lacinia mediana carnosa; toto
labello usque ad 10 mm. longo; 7 mm. lato; columna gracill,
antice villosa, usque ad 10 mm. longa; ovario pedicellato satis
gracill, usque ad 10 mm. longo.

Colombia: Dept. El Valle, Rio Calima, Quebrada de La Brea,
30-40 m. alt., SCHULTES & VILLARREAL 7370! (AMES), type!

Psilochilus physurifolius (Rchb. f.) Lgjtnant in Bot. Notiser
130: 168, 1977.
Basionym: Pogonia physurifolia Rchb.f. in Nederl. Kruidk.
4: 324, 1858.
Type: British Guiana, without precise locality, SCHOM-
BURGK s.n.! (K).

Through the courtesy of Dr. Peter Taylor, Curator of the
Orchid Herbarium at Kew, I was able to examine the type-spe-
cimen of Pogonia physurifolia. The dissection of one of the
glued-down flowers revealed a triandrous condition of the col-
umn. As a matter of fact, all details of the flower agree remark-
ably with those collected by BROADWAY s.n.! in Grenada, the
latter which was identified and illustrated by Ames as Psiloc hi-
lus macrophyllus (Lindl.) Ames. — See Ames, Orchidaceae 7:
45, Pl. 110, 1922. — The lip in P. macrophyllus differs in its
different proportions and its short claw; P. Maderoi (Schltr.)
Schltr. has a distinct keel on the long claw. The Ecuadorian
material published as P. physurifolius is based on a wrong
determination; its correct disposition will be given in the cur-
rently published Orchids of Ecuador.

Triphora Ravenii (L.O. Wms.) Garay, comb. nov.
Basionym: Pogonia Ravenii L. O. Wms. in Fieldiana, Bot.
32: 200, 1970.
Type: Costa Rica; 6 km. S. of San Vito de Java, RAVEN
21837! (F).

An examination of the holotype clearly shows that this
species is a member of the genus Triphora and closely related
to T. amazonica Schltr.

Pseudocentrum Purdii Garay, sp. nov.

Terrestris, erecta, usque ad 65 cm. alta; radicibus fascicula-
tis, carnosis, pilosis; foliis basilaribus, rosulatis, longe
petiolatis, petiolis canaliculatis, usque ad 13 cm. longis, laminis
oblique ellipticis vel ovato-ellipticis, abrupte acuminatis, her-
baceis, usque ad 18 cm. longis, 6.5 cm. latis; scapo erecto,
vaginato, supra racemoso; racemo cylindrico, satis dense mul-
tifloro, usque ad 16 cm. longo; bracteis ovato-lanceolatis,
acuminatis, 1.5 cm. longis, 0.5 cm. latis; floribus extus villosis;
sepalo postico ovato, subacuminato, usque ad 6 mm. longo, 2
mm. lato; sepalis lateralibus connatis, apice liberis, oblique
ovato-falcatis, 4 mm. longis, 3 mm. latis, deinde cylindraceo-
canaliculatis, usque ad 2 cm. longis; petalis linearibus, acumin-
atis, usque ad 4 mm. longis, | mm. latis; labello sessili, 3-lobo,
lobis lateralibus linearibus, 3 mm. longis, parallelis, deinde
abrupte in lobum intermedium, conduplicatum, linearem,
apice inflatum, decurrentibus, infra apicem lobulo fornicato
donato, toto labello usque ad 18 mm. longo; ovario gracili,
cylindrico, satis dense villoso, usque ad 11 mm. longo.

Colombia: Dept. de la Guajira, Mont del Agua, Molina, Rio
Hacha, Hacha, PuURDIE s.n.! (K).

Sobralia Fenzliana Rchb.f. in Bot. Zeit. 10: 714, 1852.
Type: Panama; Chiriqui, WARSCEWICZ 48! (W).
Syn.: Sobralia panamensis Schltr. in Fedde, Rep. Beih. 17:
11, 1922.
Type: Panama; Gatun Lake, POWELL 21! (AMES)

At the time Schlechter described S. Powellii, the type of S.
Fenzliana was not available to him, judging from an ex char.
determination of a drawing he sent to Professor Ames. An
examination of the holotype of both species confirms their
conspecificity. Recently I have seen material of this species
also from Colombia and Ecuador.

4

Sobralia luteola Rolfe in Kew Bull. 199, 1898.
Type: Tropical America: flowered in cultivation by RALLI
S.C).
Syn.: Sobralia pleiantha Schltr. in Fedde, Rep. 3: 79, 1906.
Type: Costa Rica: Boruca, PITTIER 3855! (AMES).

The rather short, distichous inflorescence with imbricating
bracts and the production of flowers in a rapid succession are
the distinguishing features of this species. An examination of
both holotypes confirms them to be conspecific.

Sobralia Powellii Schltr. in Fedde, Rep. Beih. 17: 11, 1922.
Type: Panama: near Gatun Lake, POWELL 2! (AMES).

Sobralia Powellii has been treated as a synonym of S. leu-
coxantha Rchb.f. by all contemporary students of American
orchids. An examination of the type of §. leucoxantha, how-
ever, Clearly indicates that both species are amply distinct, not
only in vegetative aspects, but also in size and structure of the
flowers. The original mix-up is most probably due to the mis-
identification of Plate 7058 in the BOTANICAL MAGAZINE. Iron-
ically, most of the true §. leucoxantha specimens in herbaria
are misidentified as §. macra Schltr., which, however, is not a
synonym of S. leucoxantha. | have seen material also from
Costa Rica, Colombia and Ecuador.

Sobralia Warscewiczii Rchb.f. in Bot. Zeit. 10: 714, 1852.

Type: Panama; Chiriqui, WARSCEWICz 8! (W).

Syn.: Sobralia Amparoae Schltr. in Fedde, Rep. Beih. 19:
8, 1923.

Type: Costa Rica; from cultivation, TONDUz 51! (AMES).
Sobralia Hawkesii Heller in Phytologia 14: 16, 1966.

Type: Nicaragua; Dept. Granada, Volcan Mombacho,
HELLER & HAWKES 30235 (FP).

An examination of the types of S. Warscewiczii and §. Am-
paroae and a comparison of them with the published illustra-
tion of S. Hawkesii indicates that all three species are con-
specific with one another.

Elleanthus Presl, Rel. Haenk. 1: 97, 1827.

Sepals free, rarely spreading; dorsal sepal concave; lateral
sepals basally connivent, gibbose. Petals free, usually thinner
in texture. Lip longer than sepals, concave with a flared apex,
basally cymbiform, excavate or gibbous with one or two dis-
tinct corpuscles or calli. Column erect, footless but with ob-
lique base, winged above. Stigmata confluent into one, promi-
nent under the triangular rostellum. Anther incumbent. Pol-
linia 8, in unequal massulae. Erect or branching shrublike
plants with fibrous roots. Stem erect, ascending or decumbent,
cane-like, leafy. Leaves distichous, rigid, plicate, sessile on
arpophyllaceous, adpressed leaf-sheath. Inflorescence termi-
nal, racemose or subcapitate, commonly rather dense and
many-flowered, rarely lax and few-flowered. Bracts conspicu-
ous, but not foliaceous. Flowers rather small, but attractively
colored.

Lectotype: Elleanthus lancifolius Pres| [Britton & Wilson
in Sci. Surv. Porto Rico 5, pt. 2: 203, 1924].

Approximately 70 species in tropical America and West
Indies.

KEY TO SECTIONS

1. Rachis shortened into a tightly packed, subglobose head; column witha
prominent, infrastigmatic protuberance forming a distinct men-
FORE ne ces area aoa oe Hew eee eae eee OSEAN Sect. Cephalelyna
la. Rachis elongate, densely or loosely spicate; column without an infras-

tigmatic mentum, at most angular-protuberant .................- 2
2. Intlorescence AtCral 665445262440s seen ndsoes ce es Sect. Laterales
Das. WMOTeSCenCe TETTINA :ecncscnectio das age sth Vea eee ee ees 3
3 Flowers and bracts in an all-sided spike or secund .............. 4

3a. Flowers and bracts in a distichous to bifarious spike, either approxi-
mate, imbricate or remote ona flexuous to strongly fractiflex raceme . 7

4. Calli of lip freely exposed in basal cavity ............. 00. e eee 5
4a. Calliof lip partially hidden behind a transverse membrane covering the
entrance to basal cavity in part ............... Sect. Hymenophora

5. Calliof lip tightly approximate or confluent in fresh condition, variously
sulcate in middle, especially when dried, centrally affixed to didymous

CAV a inen anew eas os eaten oop b Ge tL oeee CEST OeR Sect. Calelyna
Sa. Calli of lip free from one another and distinctly separated, each located
either at the center or at the base of their respective cavities ..... 6
6. Column with a biauriculate clinandrium; lateral sepals more or less
CONMMEMNG cosas ces coe eer lade cee Raa gees Sect. Otiophora

6

6a.

Ta.

Column with an exauriculate clinandrium; lateral sepals free from one
PRGRROE cna es aes eevee cee cera iro paeete Sect. Stachydelyna
Base of lip subglobose, didymous-saccate; calli inserted basally or
COMEFAIV 18 CAVITY: fae es dino cytes wees Sect. Chloidelyna
Base of lip conical, subcalcarate-saccate, not didymous; calli inserted
ON SIGS OF CAVITY 56. ieee baciee see teed rede es Sect. Elleanthus

Elleanthus Sect. Cephalelyna (Rchb.f.) Rchb.f. in Walp. Ann.

6:

474, 1862.
Basionym: Evelyna Sect. Cephalelyna Rchb.f. in Bot. Zeit.

10: 709, 1852.

Lectotype: Evelyna casapensis Rchb.f. in hoc loco.

Inflorescence terminal, capitate; rachis shortened into a

tightly packed, subglobose head; column with a prominent,
infrastigmatic mentum.

This section is the most difficult as well as the most misun-

derstood in the genus due to the superficial resemblance of its
members to one another. Since I did not find a correct treat-
ment in any of the floristic works pertaining to this group of
plants, a short synopsis with key to all species is provided here:

-

a

Y

an

wn
fet)

6a.

oa eee

KEY TO SPECIES

Lip ovate; petals linear-filiform, |-nerved; infrastigmatic protuberance
of column with a median, recurved digit ............... E. Killipii
Lip cuneate-flabellate to more or less orbicular; petals wider, several-
nerved; infrastigmatic protuberance of column without a median, re-

UNE OS SEI: ccd ds cso cng see eee eae ees tds we ae reeks 2
Lip from a concave or saccate base cuneate-flabellate ........... 3
Lip from a concave or saccate base orbicular to suborbicular .... 7
Infrastigmatic protuberance papillose-hirsute ................0.. 4
Infrastigmatic protuberance glabrous .......... 0000 c eee ee eee 6
Lateral sepals oblong-ligulate, acute; petals entire ..............5
Lateral sepals obliquely ovate-lanceolate, acuminate, petals erose-
ri oad [oa | (|| a a a ee a ee E. Glomera

Stem slender; bracts ovate; lip from a saccate base transversely
elliptic-flabellate with erose-dentate margin, retuse in front .......

OL OR cde Oh EER Ke EHR Loe NESE Re eee Chee MRE eRe e E. capitatus
Stem stout; bracts elliptic; lip from a concave base obovate-flabellate
with entire margin, bilobed in front. .................-. E. Sodiroi
Sepals more or less connate at base; lip deeply excised in middle ..
Ce eee eee Tee TTT ee or reese ee ee eee ee E. Hookeranus
Sepals free to base; lip slightly retuse in front .. E. sphaerocephalus

7

7. Infrastigmatic protuberance short, poorly developed, obscurely papil-

lose; bracts lanceolate, long-acuminate ......... E. cynarocephalus
7a. Infrastigmatic protuberance elongate, well-developed, glabrous; bracts
ovate-lanceolate, SHOrmly-ACUMINGLe- 62 cans ee Sidee es nheenewetes 8
8. Petals as wide as the sepals; lip emarginate in front with a median
GOICUM: hoy fee Stee arses dee teee rihanna ess kae eres E. cephalotus
8a. Petals narrower than sepals; lip emarginate to bilobed in front without a
PIPOUIN-ADICUIC: ooh enadcuscecscsc4e oSneet Pra eR eEek es cee eee ee 9
9. Lateral sepals 3-nerved; lip coarsely lacerate-fimbriate to lacerate-
Getitate along Margin: <4 .4.6c5 5450 oh de eee 4 aoa E. brasiliensis
9a. Lateral sepals 5- to 7-nerved; lip minutely erose-denticulate along mar-
MA. Hinges eas cae oe a8 ee ene eee rs E. casapensis

ENUMERATION OF SPECIES

Elleanthus brasiliensis (Lindl.) Rchb.f. in Walp. Ann. 6: 478,
1862.
Basionym: Evelyna Miecaieeee Lindl. in Hook., Lond.
Journ. Bot. 2: 661, 1843.
Type: Brazil; Organ einen GARDNER 642 (K).

The specimen in the Lindley herbarium is collected by
GARDNER 673! (K-L) which I examined. It is at variance with
Lindley’s drawing on the same sheet where the lip is shown as
having broadly triangular apiculate apex. However, Lindley at
the time of original publication did not describe any floral
details.

Elleanthus capitatus (Poepp. & Endl.) Rchb.f. in Walp. Ann. 6:
475, 1862.
Basionym: Evelyna capitata Poepp. & Endl., Nov. Gen. ac
Sp. Pl. 1: 32, 1836.
Type: Peru; near Cuchero, POEPPIG s.n.! (W).
Syn.: Evelyna cephalophora Rchb.f. in Bot. Zeit. 10: 709,
1852.
Elleanthus cephalophorus (Rchb.f.) Rchb.f. in Walp.
Ann. 6: 476, 1862.
Type: Peru; near Cuchero, POEPPIG 1638! (W).

The very slender plants have rather small heads. Elleanthus
cephalophorus on one of the herbarium sheets has been anno-
tated by Reichenbach himself in 1885 to be conspecific with E.
capitatus.

8

Elleanthus casapensis (Rchb.f.) Rchb.f. in Walp. Ann. 6: 475,
1862.
Basionym: Evelyna casapensis Rchb.f. in Bot. Zeit. 10: 709,
1852.
Type: Peru; Casapi, MATTHEWS 1891! (W).

This is the most common species in the Andes from Colom-
bia to Peru. The leaves are abruptly constricted below the apex
into a long-acuminate tip.

Elleanthus cephalotus Garay & Sweet inJourn. Arnold Arb. 53:
390, 1972.
Basionym: Bletia capitata R. Br. in Aiton, Hort. Kew. ed.
2, 5: 206, 1813.
Syn.: Elleanthus capitatus (R. Br.) Rchb.f. ex Cogn. in
Urban, Symb. Antill. 6: 561, 1910, not (Poepp. &
Endl.) Rchb.f., 1862.
Type: West Indies (probably Jamaica), without precise loc-
ality. Introduced by Sir JOSEPH BANKS s.n.! in 1795
and flowered at Kew Gardens Aug. 15, 1796! (BM).

This species is apparently restricted to the West Indian Is-
lands. It may be distinguished from all other members in this
section by the broad petals which are of the same width as the
sepals. A good illustration as well as a description is given in
the Orchids of Jamaica by Fawcett and Rendle.

Elleanthus cynarocephalus (Rchb.f.) Rchb.f., Walp. Ann. 6:

476, 1862.
Basionym: Evelyna cynarocephala Rchb.f., Bonpl. 4: 216,
1856.
Syn: Epidendrum capitatum Sessé & Mocino, Fl. Mex., ed.
2, 202, 1894.
Type: Mexico, Mt. Tuxtla, SEssE & MocINo s.n.! (Herb.
Pavon) (G).

Terrestrial plants, up to 50 cm. tall. Stem erect, leafy, com-
pletely enclosed by striate leaf-sheaths. Leaves elliptic-
lanceolate, long-acuminate, ribbed, up to 18 cm. long and 3.5
cm. wide. Inflorescence terminal, sessile, capitate. Bracts
congested, ovate-lanceolate to lanceolate, acuminate, up to 6

2

cm. long, decreasing inwards. Flowers rose-colored. Dorsal
sepal oblong-ligulate, obtuse, 3-nerved, up to 11 mm. long and
3 mm. wide; lateral sepals connate for 3 mm. at base, together
cymbiform; free portion linear-oblong, obliquely acute, each
up to Il mm. long and 3 mm. wide. Petals linear-oblong,
rounded at apex, erose-denticulate above, 3-nerved, up to 10
mm. long and 2 mm. wide. Lip suborbicular, lightly excised in
front with wavy and obscurely denticulate margin, slightly
gibbose at base, provided with a pair of small ovoid calli, up to
12 mm. long and wide. Column winged, more or less sigmoid
with a prominent chin. Ovary cylindric, ribbed, sessile, up to
15 mm. long.

Since Reichenbach did not describe the floral details a new
description is given here, prepared from the type. There are 3
specimens in existence. The type is in Geneva, a duplicate of it
in the Reichenbach Herbarium in Vienna, while the third, with
the original handwritten label, once part of the Lambert Her-
barium, is in the British Museum of Natural History. A good
illustration is to be found in Hamer, Las Orquideas de El
Salvador 1: 157, 1974, where it is called Elleanthus cephalotus.
The basally connate sepals are unique in the section.

Elleanthus Glomera Garay, nom. nov.
Basionym: Glomera brasiliensis Barb. Rodr., Gen. et Sp.
Orch. Nov. 1: 147, 1877.
Type: Brazil; Prov. S. Paulo, near Santos, MOSEN 3485 (S).

An examination of Rodrigues’s original drawings of floral
details of Glomera brasiliensis, now in the Library of the
Orchid Herbarium of Oakes Ames, has convinced me that this
species is sufficiently distinct from the FE. brasiliensis.

Elleanthus Hookeranus (Barb. Rodr.) Garay, comb. nov.
Basionym: Evelyna Hookerana Barb. Rodr., Gen. et Sp.
Orch. Nov. 2: 166, 1881.
Syn.: Elleanthus brasiliensis var. Hookeranus (Barb.
Rodr.) Cogn. in Mart., Fl. Bras. 3(5): 327, 1901.
Lectotype: Brazil; Prov. Rio de Janeiro, Serra d’ Estrella,
BARBOSA RODRIGUES 5.2. No specimen known

10

to exist; Rodrigues’s original drawing in the Li-
brary of the Orchid Herbarium of Oakes Ames Is
selected here for holotype! (AMES)

Elleanthus Killipii Garay, sp. nov.

Plantae elatae, usque ad 80 cm. altae; radicibus fasciculatis,
flexuosis, carnosis; caulibus erectis, primum vaginatis deinde
multifoliatis; foliis lanceolatis, acuminatis, base cuneatis, sub-
tus 5-7 nerviis, usque ad 23 cm. longis, 4.cm. latis; inflorescen-
tia terminali, capitata, dense pluriflora, ca. 2.5 cm. diametienti;
bracteis ovato-lanceolatis, acuminatis, usque ad 2 cm. longis;
floribus pallide purpureis; sepalo postico oblongo-ligulato,
acuto, basin extus sparse furfuraceo, usque ad 11.5 mm. longo,
3 mm. lato; sepalis lateralibus oblique oblongo-ligulatis, acutis
vel subacuminatis, basi excavatis, extus sparse furfuraceis,
usque ad 13 mm. longis, 3 mm. latis; petalis lineari-filiformibus,
acuminatis, uninerviis, usque ad I1 mm. longis, | mm. latis;
labello concavo, basi saccato, intus callis vel corpusculis 2,
Ovoideis, antice evanescentibus ornato, deinde in laminam
Ovatam, antice tridenticulatam expanso, margine minutissime
erosulo, toto labello usque ad 12 mm. longo, 10 mm. lato;
columna suberecta, tabula infrastigmatica 3-lobo intermedio
pubescenti, recurva, usque ad 7 mm. longa; ovario cylindrico,
pubescenti, usque ad 9 mm. longo.

Colombia: Dept. El] Valle, Buenaventura, coastal thickets,
10 m. alt., KILLIp 11760! type! (AMES), KiLLip 5215! (AMES,
US).

Elleanthus Killipii is readily distinguishable from the other
species in the Section Cephalelyna by the shape of the ovate lip
and the linear-filiform, l1-nerved petals.

Elleanthus Sodiroi Schltr. in Fedde, Rep. 14: 387, 1916.
Type: Ecuador; Pichincha near Quito, SopiRO 38! (BR).

So far this species is known only from Ecuador. An outstand-
ing field character is the entire margin to the obovate-flabellate

lip.
11

Elleanthus sphaerocephalus Schltr. in Fedde, Rep. Beih. 27: 17,
1924.
Type: Colombia; Dept. Narino, Pasto, Hopp 33.

This species is now known from Venezuela to Bolivia. As a
matter of fact, it is illustrated in Dunsterville & Garay, VENE-
ZUELAN ORCHIDS 4: 70, 1966 under the name of E. capitatus.

Elleanthus Sect. Laterales Garay sect. nov.
Type: Elleanthus lateralis Garay (in press).

Inflorescentia lateralis, spicata; columna sub fovea stigmati-
ca ut plurimum angulata, nunquam mentum formantia.

Elleanthus Sect. Hymenophora Garay, sect. nov.
Type: Evelyna hymenophora Rchb.f.

Inflorescentia terminalis, spicata; flores quaquaversae; col-
umna sub fovea stigmatica angulata; calli labelli pro maxima
parte post membranum foveae inclusi.

Elleanthus Sect. Calelyna (Rchb.f.) Benth. & Hook., Gen. PI.
3: 522, 1883.
Basionym: Evelyna Subg. Calelyna Rchb.f. in Bot. Zeit. 10:
708, 1852.
Syn.: Elleanthus Subg. Calelyna (Rchb.f.) Rchb.f. in Walp.
Ann 6: 473, 1862.
Lectotype: Evelyna myrosmatis Rchb.f. in hoc loco.

Inflorescence terminal, spicate; flowers quaquaversal; co-
lumn angulate; calli of lip freely exposed in basal cavity, tightly
approximate or confluent in fresh condition, variously sulcate
in middle when dry.

Elleanthus Sect. Otiophora Garay, sect. nov.

Type: Elleanthus Caroli Schltr.

Inflorescentia terminalis, spicata; flores quaquaversae;
sepala lateralia plus minusve connata; columna sub fovea stig-
matica angulata; clinandrium biauriculatum.

12

Elleanthus Sect. Stachydelyna (Rchb.f.) Rchb.f. in Walp. Ann.
6: 476, 1862.
Basionym: Evelyna Sect. Stachydelyna Rchb.f. in Bot. Zeit
10: 709, 1852.
Lectotype: Evelyna Lindenii Rchb.f. in hoc loco.

Inflorescence terminal, spicate; lateral sepals free; flowers
all-sided; column angulate; clinandrium exauriculate.

Elleanthus Sect. Chloidelyna (Rchb.f.) Garay, comb. nov.
Basionym: Evelyna Subsect. Chloidelyna Rchb.f. in Bot.
Zen, 102/09, 1852.
Syn.: Elleanthus Subsect. Chloidelyna (Rchb.f.) Rchb.f. in
Walp. Ann. 6: 476, 1862.
Type: Evelyna graminifolia Poepp. & Endl.

Inflorescence terminal, spicate; flowers distichous; column
angulate; base of lip subglobose, didymous, saccate; calli free,
inserted basally or centrally in cavity.

Elleanthus confusus Garay, sp. nov.

Plantae fruticosae, ramosae, supra metrales; caulibus erec-
tis, strictis, infra vaginatis, supra paucifoliatis; ramis ab-
breviatis, supra 3-, 4-foliatis, usque ad 12 cm longis; foliis
coriaceis, subplicatis, lanceolatis, subacuminatis, usque ad 7
cm. longis, 1.5 cm. latis; inflorescentia terminali, sessili, laxe
pauciflora, usque ad 3.5 cm. longa; rhachide sinuoso-
fractiflexa; bracteis ovato-cucullatis, acutis, rigidis, lepidotis,
usque ad 12 mm. longis; floribus satis tenuibus, albidis, pallide
roseo-suffusis, glabris; sepalo postico anguste ovato, concave
obtuso, plus minusve mucronato, usque ad 8 mm. longo, 3 mm.
lato; sepalis lateralibus oblique ovato-oblongis, concavis,
acutis, apice dorsaliter carinatis, usque ad 8.5 mm. longis, 3.5
mm. latis; petalis cuneato-spathulatis, obtusis, margine sub
apice utrinque denticulatis, usque ad 9 mm. longis, 4mm. latis;
labello in circuitu subrotundo, basi globoso-saccato, intus cal-
lis 2, parallelis ornato, lamina antice biloba, margine irregulari-
ter lacerato-dentata, disco ante saccum transverse incrassato;
toto labello usque ad 10 mm. longo latoque; columna arcuato-

13

clavata, alata, juxta rostellum auriculata, usque ad 5 mm.
longa; ovario cylindrico, furfuraceo.

Venezuela: Epo. ARAGUA; Rancho Grande, DUNSTER-
VILLE 533! — Ebo. BOLIVAR; Cerro Venamo, STEYERMARK &
DUNSTERVILLE 92507! Type! (AMES); Chimanta Massif,
STEYERMARK 75735! (AMES, F, NY).

This new species was illustrated in Venezuelan Orchids 2:
100, 1961 as E. virgatus (Rchb.f.) C. Schweinf., and in Lasser,
Flora de Venezuela 15(1): 226, 1970 as E. kermesinus (Lindl.)
Rchb.f. It differs from the former in the coriaceous bracts and
the differently proportioned lip; from the latter in dissimilar
floral segments and the shape of the column.

Elleanthus Sect. Elleanthus
Lectotype: Elleanthus lancifolius Presl
Syn.: Elleanthus Sect. Virgatae Garay in Orquideologia 4:
15, 1969.
Type: Sertifera virgata Rchb.f.

Inflorescence terminal, spicate; flowers distichous; column
angulate; base of lip conical, subcalcarate-saccate, not didym-
ous, Calli inserted on sides of cavity.

Elleanthus formosus Garay, sp. nov.

Plantae, erectae, verosimiliter ramosae, partes quae adsunt
usque ad 15 cm. longae; caulibus erectis, infra vagiiatis, supra
2- 4-foliatis; foliis satis coriaceis, plicatis, auguste lanceolato-
ellipticis, subacuminatis, basin cuneatis, usque ad 10 cm. lon-
gis, 1.5 cm. latis; inflorescentia terminali, sessili, usque ad 7
cm. longa; rhachide filiformi, valde fractiflexa, laxe 4-, 6-flora;
bracteis cymbiformibus, scariosis, usque ad 2 cm. longis, sur-
sum decrescentibus; floribus illis E. virgatis semillimis sed
multo majoribus; sepalo postico elliptico, apice rotundato, in
medio breviter mucronato, usque ad 11 mm. longo, 4mm. lato;
sepalis lateralibus oblique ovatis, rotundatis, breviter mucro-
natis, usque ad 12 mm. longis, 5 mm. latis; petalis e cuneata
basi obovatis apice, rotundatis, usque ad 11 mm. longis, 4mm.
latis; labello saccato, intus callis vel corpusculis lateraliter

14

insertis Ornato, lamina in ambitu suborbiculari, antice valde
biloba, margine crenulata, disco longitudinaliter incrassato in
medio, toto labello 17 mm. longo, 12 mm. lato; columna cylin-
drica, usque ad 7 mm. longa; ovario cylindrico, obscure alato,
usque ad 7 mm. longo.

Colombia: DEPT. NARINO, on road Barbacoas-Tuqueres,
SCHMIDTCHEN s.n.! Type! (W).

Cybebus Garay, gen. nov.
Etymology: Greek Kybebos = stooping, with head bent, in
reference to the rectangularly bent flowers.

Sepala inter se breviter in tubum cylindricum, arcuatum
connata, deinde libera, arcuato-patentia. Petala sepalo postico
conniventia. Labellum horizontale, naviculare. Columna re-
ctangulariter arcuata, dimidio superiore porrecta, libera, di-
midio inferiore sepalis connata, et cum eis pedem elongatum,
arcuatum formans; clinandrium amplum; rostellum transver-
sum, 3-lobum, lobo intermedio lineari-triangulo; stigmata 2,
sub rostello confluentia.

Herbae terrestres elatae, grandiflorae; radicibus fasciculatis
tuberosis; foliis basilaribus, plurimis, petiolatis; scapo erecto,
Vaginato, supra laxe paucifloro; bracteis prominentibus,
ovariis longioribus; floribus speciosis, magnis; Ovario cylin-
drico, sessill.

Type: Cybebus grandis Garay

Species singula, adhuc nota, Colombiana.

This new genus is closest to Mesadenus Schltr. in the struc-
ture of the rostellum and in the general appearance of the
flowers. The rectangularly arcuate column with a long foot,
and the tubulose base of the sepals readily differentiate this
genus from the other members of the Spiranthes alliance.

Cybebus grandis Garay, sp. nov.

Terrestris, erecta, usque ad 50 cm. alta; radicibus fas-
ciculatis, tuberosis, villosis; foliis basilaribus, plurimis, in
specimine nostro 5-nis, petiolatis; petiolis canaliculatis, basin
imbricantibus, usque ad 10 cm. longis; laminis oblique ovato-
ellipticis, acutis, basi acuto-cuneatis, usque ad 12 cm. longis,

15

5.5 cm. latis; scapo erecto, plurivaginato, infra glabro, supra
puberulo, tertia parte superiori laxe paucifloro; bracteis linea-
ri-lanceolatis, acuminatis, Ovariis duplo superantibus, usque
ad 5.5 cm. longis; floribus speciosis, extus puberulis; sepalis
inter se in tubum arcuato-cylindricum breviter connatis, de-
inde liberis, arcuato-patentibus; sepalo postico anguste
lanceolato-elliptico, acuminato, usque ad 35 + 5 mm. longo, 7
mm. lato; sepalis lateralibus lineari-lanceolatis, acuminatis,
usque ad 35 + 5mm. longis, 5 mm. latis; petalis e cuneata basi
falcato-dolabriformibus, acuminatis, omnino glabris, usque ad
35 + 5 mm. longis, 7 mm. latis; labello porrecto, naviculari,
apice recurvo, basi biauriculato-sagittato, usque ad 35 mm.
longo, 10 mm. lato; columna arcuata, facie puberula, usque ad
18 mm. longa; rostello prominenti, 3-lobo, 2 mm. longo, 3 mm.
lato; ovario cylindrico, puberulo, torto, usque ad 3.5 cm.
longo.

Colombia: Depto. del Cauca, near lake El Tambo, Munchi-
que, K. VON SNEIDERN 731! type! (S).

Cyclopogon Presl, Rel. Haenk. |: 93, t.13, f-1, 1827.
Etymology: Kyklos = circle and pogon = tail [1.e., of fire],
with divided ends, in reference to the (reddish?)
sepals which arise in a circle from a tube, like
tails of fire with divided ends.
Type: Cyclopogon ovalifolium Presl.

The diagnostic features of the genus Cyclopogon for the past
125 years have been completely misunderstood. Cyclopogon
was generally regarded a synonym of Spiranthes since 1840,
when Lindley transferred it in his Genera and Species of Or-
chidaceous Plants. In 1920, Schlechter reinstated it in his study
of the Spiranthianae, with a rather detailed generic circum-
scription. Unfortunately both of these students of orchidology
overlooked the fact that Haenke’s illustrations show a peculiar
character, albeit not mentioned in the original description,
namely, the sepals are connate with one another to form a
distinct sepaline tube.

Recently I have seen several collections from Ecuador and
Peru referable to C. ovalifolium which | intended to describe as

16

anew genus, based on the fusion of the sepals. Once I became
aware of the characters of the true Cyclopogon, I requested the
type from Prague, and an examination of it confirmed my
interpretation. Cyclopogon is closest to the genus Gamosepal-
um, but in the latter genus the column in the flowers is adnate
dorsally to the sepals.

It must be emphasized that the circumscription given by
Schlechter for Cyclopogon is no longer tenable. Fortunately,
for Schlechter’s concept the generic name, Beadlea Small is
available. Appropriate transfers are being made in the new
generic revision of the Spiranthinae by Dr. Pabst and me.

The following amplified descriptions are prepared from the
type-material and from the additional material at hand. The
genus is monotypic.

Sepals similar, basally connate into a cylindric tube, above
free. Petals connivent with dorsal sepal, basally for a short
distance adnate to column. Lip broadly unguiculate, sagittate-
auriculate, the claw adnate to sepaline tube. Column free,
elongate, slender, cylindric, pubescent in front, basally pro-
duced into a short, somewhat descending foot; stigmata 2,
approximately touching each other; rostellum elongate, ligu-
late, truncate or obscurely excised; anther in the descending
clinandrium dorsal, erect, 2-celled. Ovary sessile. Terrestrial
plants with fleshy, fasciculate, villose roots. Leaves basal,
many, petiolate. Scape slender, several-sheathed, spicate
above. Bracts linear-setaceous. Flowers small.

Cyclopogon ovalifolium Presl, Rel. Haenk. |: 93, 1827.
Syn.: Spiranthes Preslii Lindl., Gen. and Sp. Orch. PI. 470,

1840.
Gyrostachys ovalifolia (Presl) O. Ktze., Rev. Gen. PI.
2: 664, 1891.

Type: Peru; Prov. Huanuco, HAENKE s.n.! (PR).

Terrestrial plants, up to 35 cm. tall. Roots fasciculate, rather
thick, fleshy, villose. Leaves basal, up to 5, petiolate; blade
obliquely elliptic to ovate-elliptic, acute, basally rounded or
abruptly cuneate, up to 7.5 cm. long and 4cm. wide, commonly

smaller; petiole slender, canaliculate, up to 5 cm. long. Scape

17

erect, slender, loosely several-sheathed, terminated by a one-
sided spike; spike cylindric, subdensely many-flowered, very
variable in length, 4 to 14 cm. long. Bracts linear to linear-
setaceous, up to 15 mm. long. Flowers small, tubular, greenish
or brownish green. Sepals similar, linear-ligulate, obtuse, bas-
ally connate into a distinct, cylindrical tube which Is 2.5 to 3.5
mm. long, including the tube up to 8.5 mm. long and 1.5 mm.
wide. Petals linear-subspathulate, rounded, up to 8 mm. long
and | mm. wide. Lip unguiculate, sagittate-auriculate at base,
the auricles flat and more or less rounded; blade linear-oblong,
constricted at one-fourth from apex producing a transversely
elliptic, terminal lobe; disc pubescent; whole lip up to 8.5 mm.
long and 3 mm. wide in front. Column slender, up to 6 mm.
long. Ovary sessile, glabrous, cylindric, up to 4 mm. long.

Ecuador: IMBARBURA; along trail to Rio Chalguayaco, below
Magnolia, lower Intag Valley, DREW E-587! (AMES). —
PICHINCHA: On western side of Cerro Pichincha, JAMESON 38!
(K-L). — PASTAZA: in the vicinity of Colonia Jativa, ca. 12 km.
from Mera, H. LuGo 98! (AMES, GB). — Peru: HUANUCO,
HAENKES.n.! (PR). —SAN MARTIN, Roque, MELIN I11!(S).

Sauroglossum andinum (Hauman) Garay, sfat. nov.
Basionym: Spiranthes nitida var. andina Hauman in Anal.
Soc. Cienc. Argent. 90: 123, 1921.
Lectotype: Argentina; Prov. Catamarca, Rodeo, CASTIL-
LON 2042!, in hoc loco.

In general appearance the plants of S. andinum are very
similar to S. distans Lindl. ex Garay, but the floral segments,
especially the shape of lip, are amply distinct from one another.

Sauroglossum aurantiacum (C. Schweinf.) Garay, stat. nov.
Basionym: Spiranthes Weberbaueri var. aurantiaca C.
Schweinf. in Bot. Mus. Leaf. Harv. Univ. 15: 7,
1951.
Type: Peru; Prov. Urubamba, Piri, VARGAS 5935! (AMES).

In general appearance the plants are like those of S$. longi-
florum (Schltr.) Garay (comb. nov. in press); in floral detail
they are reminiscent of §. e/atum Lindl.

18

Sauroglossum corymbosum (Lindl.) Garay, comb. nov.
Basionym: Synassa corymobsa Lindl. in Bot. Reg. 19: sub
t. 1618, 1833.

Syn: Pelexia corymbosa (Lindl.) Lindl., Gen. and Sp. Orch.
Pl. 482, 1840.
Synassa dilatata Lindl. ex Krzl. in Engl., Bot. Jahrb.
54, Beibl. 117: 19, 1916, sphalm.
Spiranthes subumbellata C. Schweinf. in Bot. Mus.
Leafl. Harv. Univ. 10: 31, 1941.

Type: Peru; without precise locality, PAVON s.n.! (K-L).

A point-by-point comparison of the characters I have ob-
served in the dissected flowers of the type-specimens of
Synassa corymbosa Lindl. and Sauroglossum elatum Lindl.
failed to disclose any criteria by which the two species could be
kept apart generically.

Sauroglossum distans Lind]. ex Garay, sp. nov.

Terrestris, erecta, usque ad 60 cm. alta; foliis basilaribus,
ellipticis vel obovato-ellipticis, basin versus subpetiolato-
angustatis, acutis, usque ad 13 cm. longis, 3.3 cm. latis; scapo
erecto, primum vaginis imbricatis obtecto, supra laxe multi-
floro; bracteis lanceolatis, acuminatis, usque ad 2 cm. longis,
sursum decrescentibus; floribus extus pubescentibus; sepalo
postico lineari-oblongo, concavo, acuto, usque ad 10 mm. lon-
go, 2mm. lato; sepalis lateralibus oblique lineari-oblanceolatis,
acutis, usque ad 12 mm. longis, 2 mm. latis; petalis e lineari basi
oblanceolatis, acutis, basi decurrentibus, usque ad 10 mm.
longis, 1.2 mm. latis; labello e linari basi spathulato, lamina
plus minusve rhomboidea, margine subcrenulata, disco basin
obscure bicalloso, in medio laminae linea incrassata ornato;
toto labello usque ad 13 mm. longo, 4 mm. lato; columna
gracilil, glabra, usque ad 8 mm. longa; ovario fusiform1,
puberulo, brevissime pedicellato, usque ad 10 mm. longo. —
Sauroglossum distans Lindl. nomen nudum in Ann. Mag. Nat.
Hist. ser. 3, 1: 334, 1856.

Bolivia: without precise locality, BRIDGES s.n./ type! (K).

19

Hapalorchis lineatus (Lindl.) Schltr. in Beih. Bot. Centralbl.
37(2): 363, 1920.
Basionym: Spiranthes lineata Lindl., Gen. and Sp. Orch.
Pl. 471, 1840.
Syn.: Gyrostachys lineata (Lindl.) O. Ktze., Rev. Gen. PI.

2: 664, 1891.
Cyclopogon lineatus (Lindl.) Pabst in Bradea |: 466,
1975.
Lectotype: Brazil; Sao Paulo, Martius s.n.! (M), in hoc
loco.

Sauroglossum monophyllum Griseb., Symb. Fl. Ar-
gent. 339, 1879.

Type: Argentina; Prov. Oran, LORENTZ & HIERONYMUS
286 (CORD).
Sauroglossum tenue Lindl. in Ann. Mag. Nat. Hist.
ser. 3, 1: 334, 1856.
Spiranthes tenuis (Lindl.) Benth. ex Fawc., FI. Pl.
Jam. 40, 1893, not Lindl. 1840.
Spiranthes Fawcettii Cogn. in Fedde. Rep. 7: 123,
1909.
Hapalorchis tenuis (Lindl.) Schltr. in Beih. Bot. Cen-
tralbl. 37 (2): 363, 1920.

Type: Cuba; Monte Verde, WRIGHT 622! (K-L).
Spiranthes amabilis Ames, Sched. Orch. 2: 8, 1923.

Type: Guatamala; Dept. Alta Verapaz, Coban, TUERCK-
HEIM II-1787! (US).
Cyclopogon stenoglossus Pabst in Bradea I: 468,
1975.

Type: Brazil; Minas Gerais, near Caxambu, WELTER 188!
(HB).

While reviewing the genus Hapalorchis for the Flora of
Ecuador, it became apparent that Lindley included two dif-
ferent species under the original description of Spiranthes
lineata. He examined Martius’ collection which is mostly in
fruit and included with it a drawing by Descourtilz prepared in
the field. Since a specimen always takes precedence over draw-
ings when available, the Martius collection must be regarded as
the lectotype for the species. I have studied this collection and

20

it shows an autogamous plant in which the pollinia wrap them-
selves around the rostellum. Consequently, when the flowers
are dissected, the rostellum always becomes ruptured and
torn, showing a 3-dentate clinandrium. The situation is the
same in the type-collections of Sauroglossum tenue, Spiran-
thes amabilis and Cyclopogon stenoglossus. It is rather obvi-
ous that we are dealing with a rather widespread species with
very little internal variation. On the other hand, the Descourtilz
drawing represents a distinct, outcrossing species which is
limited in distribution to southern Brazil. For those plants I
propose the name Hapalorchis Lindleyanus.

Hapalorchis Lindleyanus Garay, sp. nov.

Terrestris erecta, usque ad 30 cm. alta; radicibus crassis,
villosis; foliis plurimis, petiolatis, petiolis canaliculatis,
gracilibus, base vaginatis, usque ad 3 cm. longis; laminis
Ovatis, acutis, basi rotundatis, usque ad 4.5 cm. longis, 2 cm.
latis; pedunculo erecto, vaginifero, supra laxe paucifloro; brac-
teis ovato-cucullatis, longe acuminatis, usque ad 1.5 cm. lon-
gis, sursum decrescentibus; floribus satis tenuibus, extus basin
sparse pubescentibus, secundis; sepalo postico lineart-
oblongo, concavo, apiculato, usque ad 8 mm. longo, 2 mm.
lato; sepalis lateralibus porrectis, oblongo-linearibus, acutis,
usque ad 8 mm. longis, 1.2 mm. latis; petalis lineari-spathulatis,
subacuminatis, margine superiore subcrenulatis, usque ad 8
mm. longis, 1.3 mm. latis; labello ima basi excavato, lamina
primum subquadrata, deinde in lobum reniformem expansa,
disco in centro pubescenti, basin margine incrassata, antice
lineis ternis valde discoloribus ornata; toto labello usque ad 7
mm. longo, 5 mm. lato; columna usque ad 5 mm. longa; ovario
ovoideo, pubescenti, usque ad 8 mm. longo.

Brazil: Therezopolis, Rio, M. & R. FOSTER 1005! type!
(AMES).

As I stated above, the specimens referable to this rather
common species have always been confused with H. lineatus
from which it differs in having larger floral segments and dis-
similar lip.

|

Pelexia Hameri Garay, sp. nov.

Terrestris, gracilis, usque ad 30cm. alta; radicibus fascicula-
tis, tuberosis, villosis; foliis basilaribus, ut videtur 2, sub
anthesin jam emarcidis, ovatis, obtusis vel subacutis, distincte
breviter petiolatis, petiolis inclusis usque ad 5 cm. longis, 1.8
cm. latis; scapo gracili, plurivaginato, puberulo, supra laxe
multifloro, spicato; bracteis ovato-lanceolatis, acuminatis,
usque ad 9 mm. longis; floribus viridi-albidis, extus sparse
pubescentibus; sepalo postico anguste ovato-lanceolato vel
ovato-elliptico, obtuso, usque ad 3.8 mm. longo, | mm. lato;
sepalis lateralibus oblongo-ligulatis, obtusis, usque ad 4 mm.
longis, 1.2 mm. latis, basin oblique saccatis, sacco 2 mm.
longo; petalis sepalo postico conniventibus, e cuneata basi
oblique lineari-dolabri formibus, obtusis, usque ad 4 mm. lon-
gis, | mm. latis, labello unguiculato, basi lineari-falcato sagit-
tato, lamina conduplicata, lineari-oblonga, apice abrupte in
lobum reniformem expansa, margine repanda; disco basin lobi
terminali papilloso-incrassato; toto labello 5.5 mm. longo, 3
mm. lato; columna infundibuliformi, 2 mm. longa, basi in
pedem decurrentem, 2 mm. longam producta; ovario arcuato-
fusiformi, puberulo, sessili, usque ad 6 mm. longo.

EI Salvador: in the gardens of Dona Hertha Freund in San
Benito, San Salvador, HAMER 613! type! (AMES).

The plants are reminiscent of weak specimens of P.
Schaffneri (Rchb.f.) Schltr., but the flowers are very different.
This smallest-flowered Pelexia is dedicated to Mr. Fritz
Hamer. whose lavishly illustrated, two-volume work on the
Orchids of El Salvador is one of the most outstanding contribu-
tions to our knowledge of the orchids of Central America.

Sarcoglottis Lehmannii Garay, sp. nov.

Terrestris, elata, usque ad 30cm. alta; radicibus fasciculatis,
crassis, villosis; foliis rosulatis, petiolatis; petiolis ca-
naliculatis, usque ad 6 cm. longis, laminis late ellipticis, utrin-
que obtusis vel subacutis, usque ad 16 cm. longis, 6 cm. latis;
scapo erecto, vaginis acuminatissimis, imbricatibusque ob-
tecto, supra laxe racemoso, paucifloro; bracteis ovato-lanceo-
latis, acuminatissimis, usque ad 2.5 cm. longis; floribus, parvu-

22

lis, extus dense pubescentibus; sepalo postico anguste ellip-
tico, acuto, usque ad 12 mm. longo, 3 mm. lato; sepalis laterali-
bus ovario longe decurrentibus, oblique lanceolatis, subacu-
minatis, usque ad 10 + 18 mm. longis, 3 mm. latis; petalis e
cuneata basi oblique anguste falcato-ellipticis, acutis, margine
minute ciliolatis, usque ad 11 mm. longis, 2 mm. latis; labello
dimidio basali anguste lineari, deinde in laminam ovatam, ca-
naliculatam, intus sparse pubescentem expanso, apice lobulo
cordiformi, acuminatissimo, recurvo producto, basi auriculis
filiformibus, 4mm. longis donato, toto labello usque ad 25 mm.
longo, 4 mm. lato; rostello 2.5 mm. longo; ovario cylindrico,
pubescenti, usque ad 15 mm. longo.

Colombia: DEPT. CAUCA, Timbiqui, LEHMANN s.n.! type.
[H.K. 1263] (K).

Buchtienia rosea Garay, sp. nov.

Terrestris, elata, usque ad 120 cm. alta; radicibus fas-
ciculatis, carnosis, villosis; foliis basilaribus, ut videtur duis,
longe petiolatis, petiolis canaliculatis, robustiusculis, usque ad
29 cm. longis, laminis late ellipticis, subacutis vel obtusis, basi
subcordato-cuneatis, usque ad 23 cm. longis, 13 cm. latis;
scapo erecto, primum vaginato, deinde laxe racemoso; racemo
cylindrico, multifloro, usque ad 35 cm. longo; bracteis lineari-
lanceolatis, anguste acuminatis, usque ad 2 cm. longis, sursum
decrescentibus; floribus carnosis, roseis; sepalo postico an-
guste elliptico, acuto, extus sparse puberulo, usque ad 9 mm.
longo, 3 mm. lato; sepalis lateralibus oblique ovato-lanceolatis,
subacuminatis, extus sparse puberulis, usque ad 12 mm. lon-
gis, 3 mm. latis; petalis oblique falcato-ellipticis, utrinque at-
tenuatis, acutis, usque ad 9 mm. longis, 2 mm. latis; labello e
basi subquadrato aequaliter cuneato-trilobo, lobis lateralibus,
oblongis, apice rotundatis, lobo intermedio triangulari, obtuso,
2 mm. longo, disco basin marginaliter incrassato; toto labello 7
mm. longo, 6mm. lato; ovario cylindrico, pubescenti, usque ad
11 cm. longo.

Peru: Dept. Cuzco, on road from Mistiana to Keros, VARGAS
7381! type! (AMES).
It differs from B. boliviensis Schltr. in the color of the flow-

a2

ers, in the not long-acuminate sepals and the differently prop-
ortioned lip which is equally 3-lobed in front with a very short
midlobe.

Eltroplectris pauciflora (Poepp. & Endl.) Garay, comb. nov.
Basionym: Pelexia pauciflora Poepp. & Endl., Nov. Gen.
ac Sp. Pl. 2: 17, t. 124, 1837.

Syn.: Physurus pauciflorus (Poepp. & Endl.) Lind]l., Gen.
and Sp. Orch. Pl. 504, 1840.
Erythrodes pauciflora (Poepp. & Endl.) Ames, Or-
chid. 7: 75, 1922.

Type: Brazil; Prov. Amazonas, Rio Negro, around Ega,
POEPPIG s.n.! (W).

It is difficult to understand why this plant was treated as a
member of the genus Physurus or Erythrodes by former stu-
dents of orchidology. The plants have neither rhizome nor
cauline leaves, nor have the lips a distinct spur. An examina-
tion of the type undoubtedly places it in the genus Eltroplectris,
and its closest relative is another Brazilian species, namely, E.
longicornu (Cogn.) Pabst.

Platythelys peruviana Garay, sp. nov.

Terrestris, parvula, usque ad 12.5 cm. alta; rhizomate as-
cendenti, pluriarticulato; caulibus gracilibus, infra vaginatis,
supra laxe paucifoliatis; foliis lineari-lanceolatis, acuminatis,
basi rotundatis, breviter petiolatis, petiolo incluso usque ad 3.5
cm. longis, 0.6 cm. latis; inflorescentia terminali, abbre viata,
pauciflora, usque ad 2.2 cm. longa; pedunculo 1.5 cm. longo;
floribus satis parvulis; sepal postico anguste ovato, obtuso,
concavo, usque ad 3.5 mm. longo, |.2 mm. lato; sepalis latera-
libus oblique oblongo-ligulatis, obtusis, usque ad 3.5 mm. lon-
gis, 1.2 mm. latis; petalis Cuneato-oblanceolatis, acutis vel
subacuminatis, usque ad 3 mm. longis, 0.5 mm. latis; labello
calcarato, calcare cylindrico, usque ad 3 mm. longo, lamina
bipartita: hypochilio valde concavo, antice emarginato, epi-
chilio late cordato-reniformi, acuminato, utrinque rotundato,
toto labello 3 mm. longo, 1.5 mm. lato; ovario arcuato-cylin-
drico, glabro, usque ad 6 mm. longo.

24

Peru: Dept. Junin, Chanchamayo Valley, SCHUNCKE 569!
type! (F).

Specimens of this species have been reported in the litera-
ture erroneously as Erythrodes querceticola (Lindl.) Ames.

Stelis trigoniflora (Sw.) Garay, comb. nov.
Basionym: Epidendrum trigoniflorum Sw., Nov. Gen. &
Sp. Pl., Prodr. 125, 1788, excluding synomyms.
Syn.: Stelis ophioglossoides Auct. not Sw. in Schrad.,
Journ. Bot. 2 (4): 239,1800.

When Swartz described Epidendrum trigoniflorum, he
based his description on his own collection from Jamaica. He
also prepared detailed drawings of it for inclusion in his Icones
Plantarum Incognitarum, t. 49, but they remained unpublished.
Under Epidendrum trigoniflorum Swartz cited in synonymy
the plate of Epidendrum ophioglossoides, t. 133, f. 2, which
Jacquin published in 1763. In 1791, in his Observationes
Botanicae, p. 332, under E. ophioglossoides Swartz remarks:
**Plantae Plumieri & Jacquini certe distinctae. ... JAC-
QUINI... ad E. trigoniflorum (Prodr. p. 125) pertinet, affir-
mante planta Ipsius in Museo BANKSIANO servata.”’ I have
examined the Jacquin material and it is specifically different
from Epidendrum trigoniflorum, while the Plumier material is a
true Pleurothallis.

Although convinced that the Plumier and Jacquin illus-
trations and material were different from one another, in 1800,
in proposing the genus Sre/is, Swartz united them under Stelis
ophioglossoides under which name the error persisted until
recently (See Garay & Sweet in Journ. Arnold Arb. 53: 391 and
528, 1972). I have recently noticed that Turton has already
separated Epidendrum ophioglossoides Jacq. and Epidendrum
trigoniflorum Sw. as two distinct species in his Vegetable
Kingdom 2: 1508 and IS5I1, 1806.

Octomeria truncata Hoffmsgg., Preis. Verz. Orch. 26, 1842.
Type: Brazil; Rio de Janeiro, HOFFMANSEGG s.n.! (W).
Syn.: Octomeria lobulosa Rchb.f. in Hamb. Gartenz. 14:

215, 1858.

25

Type: Brazil; without precise locality, cult. SCHILLER s.n.!
(W).

It is difficult to understand why Reichenbach described O.
lobulosa when the actual type-material of O. truncata, both
drawings and specimens were in his possession since 1840.
Morphologically the specimens and flowers of both species are
indistinguishable.

Epidendrum tropidioides Garay, sp. nov.

Plantae procurrentes, valde ramosae; caulibus primum vag-
inantibus, supra paucifoliatis, compressis; foliis lanceolato-
oblongis, acutis vel subacuminatis, basi caulem amplecten-
tibus, usque ad 20 cm. longis, 2.2 cm. latis; inflorescentiis
terminalibus, arcuatis; pedunculo prominenti, compresso,
alato, usque ad 13 cm. longo, apice ramoso; ramis racemosis,
arcuatis, usque ad I5 cm. longis; bracteis distichis, ovato-cu-
cullatis, acutis, usque ad 1.5 cm. longis, illis Tropidiis in men-
tem revocantibus; floribus satis firmis, magnis, pallide viridis,
extus purpureo-striatis; sepalo postico anguste lanceolato,
acuto, usque ad 2.2 cm. longo, 5 mm. lato; sepalis lateralibus
dorsali simillimis sed paulo angustioribus; petalis e cuneata
basi rhombeo-spathulatis; acutis, margine interiore sublobatis,
usque ad 2.2 cm. longis, 9 mm. latis; labello in ambitu
cuneato-obovato, antice trilobo, lobis lateralibus brevibus,
rotundatis, lobo intermedio antico, suborbiculari, acuto, mar-
gine erosulo; disco supra basin bicarinato, toto labello 2.2 cm.
longo, 8 mm. lato; columna humili, clinandrio urceolato, usque
ad 8 mm. longo: ovario cylindrico, glabro, usque ad 1.5 cm.
longo.

Colombia: DEPT. PUTUMAYOo, 5 km. N.E. of Sibundoy,
BRISTOL 388! type! (AMES).

Chondrorhyncha stenioides Garay, sp. nov.

Epiphytica, caespitosa, humilis, usque ad 15 cm. alta;
radicibus carnosis, leviter flexuosis, glabris; caulibus nullis;
foliis ad 3, distichis, obovato-oblanceolatis, acutis, satis tenui-
bus, basi cuneatis et vaginis conduplicatis articulatis, usque ad
13 cm. longis, 3 cm. latis, in exsiccatione angustioribus:

26

pedunculo laterali, gracili, unifloro, usque ad 3 cm. longo;
bractea Ovato-cucullata vel infundibuliformi, usque ad 7 mm.
longa; floribus satis magnis, apertis, albidis vel viridi-albidis,
labello intus obscure purpureo-maculato; sepalo positco valde
concavo-cucullato, in ambitu subrotundo-elliptico, obtuso,
usque ad 20 mm. longo, 16 mm. lato; sepalis lateralibus pat-
entibus, ovato-ellipticis, basin versus excavatis, obtusis,
usque ad 24 mm. longis, 10 mm. latis; petalis cum sepalo
postico conniventibus, oblique late ellipticis, obtusis, usque ad
20 mm. longis, 12 mm. latis; labello valde carnoso, complicato,
in ambitu triangulo, lobis lateralibus inter se subquadratis,
replicatis, deinde in lobo terminali, rhynchophoro, tubuloso
continuis, disco a basi usque ad medium carina sigmoidea
decoro, in medio callo denticulato, transverso ornato, toto
labello usque ad 13 mm. longo, 10 mm. lato; columna cylin-
drica, usque ad 9 mm. alta, basi in pedem, 8 mm. longam
producta; polliniis 4, inaequalibus, obpyriformibus; ovario
costato, glabro, usque ad 12 mm. longo.

Ecuador: Prov. Pastaza, at confluence of Rio Verde and Rio
Pastaza, STACY s.n.! type! (AMES).

The flowers of these plants closely resemble those of Stenia
pallida, having in common the general configuration of the lip.
The method of adnation of the lip to the column-foot, the
structure of the pollinia are those of the genus Chon-
drorhyncha. This new species is perhaps closest to C. guttata
(Rchb.f.) Garay from Peru from which it differs in having larger
flowers without any spots, and differently proportioned floral
parts.

Chondrorhyncha guttata (Rchb.f.) Garay, comb. nov.
Basionym: Stenia guttata Rchb.f. in Gard. Chron. n.s. 14:
134, 1880.
Type: Peru; without precise locality, DAvis s.n.! (W).

I have seen about seven different collections of this rare
species, all from Tingo Maria, Muna and San Martin regions of
Peru. The narrower floral segments of the strongly maculate
flowers readily differentiate it from C. stenioides Garay.

oe |

Maxillaria pardalina Garay, nom. nov.
Basionym: Maxillaria pantherina Rchb.f. in Bonpl. 3: 239,
1855, not Hoffmsgg. 1844.
Type: Ecuador; Andes [of Quito] JAMESON 761! (W).

Since the specific epithet ‘‘pantherina’’ is preoccupied by a
Brazilian species, the Andean plants require a new name.

Although Maxillaria pantherina Rchb.f. has been reduced to
a synonym of M. platpetala Ruiz & Pav. by Brieger and Hunt
in 1969, their judgment is based on a single specimen in the
Lindley Herbarium. They assumed it to be an isotype of M.
platypetala, overlooking the fact that Lindley clearly stated in
his Genera and Species of Orchidaceous Plants, p. 143, 1832,
that his only specimen was donated to him by Hooker and it
was collected by Hall in Ecuador. When Reichenbach de-
scribed his M. pantherina, he was aware of Lindley’s interpre-
tation of M. platypetala, for he clearly says ‘‘Maxillariam
platypetalam nisi Ruizianam tamen Lindleyanam non novi,
quae nonnullis nobis quadrare videtur. Cum tamen cel. auctor
procurrentem dicat similliman platypetalae, haec longe certe
distat.”’

It seems advisable to repeat Hoffmannsegg’s original de-
scription for it is very inaccessible; some of its peculiar termin-
ology is explained in parentheses:

Maxillaria pantherina Hoffmsegg. — Plantae insidentes, nu-
tricariae (pseudobulbose with fasciculate roots), nutricio sterill
(leafbearing pseudobulb) pyramidali sulcato-costato, follis
binis oblongis et linearibus, inflorescentia laterali, pedunculis
(sub-) binis bracteatis, indumentis (bracts) conformibus, ob-
longis, acuminatis, sulphureo-pomaceis, petalis sepalis ¥3 bre-
vioribus dimidio angustioribus, labello 3-lobo albido, circa
marginem maculis irregulariter sparsis atropurpureis. Ibid.
(Rio de Janeiro). — Judging from the rather poor drawing by
Hoffmansegg, M. pantherina most probably is identical with
M. marginata (Lindl.) Fenzl.

Osmoglossum candidum (Linden & André) Garay, comb. nov.
Basionym: Odontoglossum candidum Linden & Andre in
Ill. Hortic. 22: 58, 1875.

28

Type: Colombia: Depr. Cauca, without precise locality,
LINDEN S5.n.

The genus Osmoglossum is new to Colombia, and plants must
be exceedingly rare, for I have not yet seen a specimen col-
lected in the wild. It appears to be close to O. Egertonii
(Lindl.) Schltr., but differs in the pandurate lip and the fim-
briate column-wings.

Osmoglossum panduratum Garay, sp. nov.

Epiphytica, usque ad 50 cm. alta; rhizomate repenti, valida;
pseudobulbis approximatis, anguste ovoideis, compressis,
2-foliatis, usque ad 9 cm. longis, 3 cm. latis; foliis oblongo-lin-
earibus, loratis, acutis, usque ad 42 cm. longis, 1.5 cm. latis;
inflorescentiis erectis vel suberectis, laxe paucifloris, usque ad
30 cm. longis; pedunculis ancipitibus, paucivaginatis; bracteis
lineari-lanceolatis pedicellis aequilongis, usque ad | cm. lon-
gis; floribus niveis, columna pallide rosea, callo aurantiaco,
brunneo-punctato; sepalo postico lanceolato-elliptico, abrupte
acuminato, 14 mm. longo, 5 mm. lato; sepalis lateralibus ter-
tiam partem basalem inter se connatis, oblique oblongo-lanceo-
latis, abrupte acuminatis, 14 mm. longis, 4 mm. latis; petalis
late ellipticis, obtusis vel subacutis, 15 mm. longis, 8 mm. latis;
labello sessili, pandurato, a medio replicato, basi truncato,
antice ovato, abrupte acuto, disco callo W-formi, valde car-
noso a basi usque ad medium ornato, explanato 13 mm. longo,
8 mm. lato; columna generis, clinandrio denticulato; ovario
cum pedicello geniculato.

Ecuador: without precise locality. Imported and grown by
JOHN STACY s.n.! type! (AMES).

This new species is closely related to O. pulchellum differing
from it in its much smaller flowers, and in the shape of the lip. In
O. pulchellum the constriction of the lip is in front of callus, the
terminal lobe is larger than the basal part, subquadrate to subro-
tund, truncate in front and the callus occupies % of the disc of
lip. In our plants the constriction of the lip is on the sides of
callus, the terminal lobe is smaller than the basal part, ovate,
abruptly subacute and the callus occupies % of the disc of the lip.

29

Miltonia flava Lind]. in Gard. Chron. 475, July 15, 1848; Baxt.

in Loudon, Hort. Brit. Suppl. 3: 589, 1850.

Type: Brazil, without precise locality, LODDIGES s.n.!
(K-L).

Syn.: Odontoglossum anceps K\. in Allgem. Gartenzeit. 19:
250, 1851;
Miltonia anceps (K\.) Lindl., Folia Orch. Miltonia 3,
1853.
Oncidium anceps (KI.) Rchb.f. in Walp. Ann. 6: 758,
1863.

Type: Brazil; without precise locality, ALLARDT s.n.

Miltonia flava has always been an enigmatic species and is
still in want of rediscovery. It is known to me only through
specimens in the Lindley and Reichenbach Herbaria. The spe-
cimen itself is a document of a strange historical background.

Introduced from Brazil by Loddiges around 1843, it flowered
but once when it was sent to Lindley for naming. Lindley
prepared a fine drawing in color from the already withering
flower which is mounted together with the specimen in his
herbarium, and annotated by his hand as Miltonia flava. At that
time Lindley must have also prepared a description, which,
however, was not published until July 15, 1848 in the GARDE-
NERS’ CHRONICLE, and then anonymously. Baxter, however,
not only reported it in Loudon’s HoRTUs BRITANNICUS Suppl.
3: 589, 1850, but also attributed the name to Lindley. Yet for
reasons unknown Lindley, as well as Reichenbach, completely
ignored the publication of Miltonia flava. In 1853, curiously
enough, while writing the pages for his FOLIA ORCHIDACEA,
Lindley annotated his specimen of Miltonia flava as Miltonia
anceps and has described it under that latter name without any
reference to the earlier publication. Since the original descrip-
tion of Milton flava is not readily available, it is repeated here:

“Sp. Char. — Flower-stalk one-flowered, with equitant cari-
nate sheaths. Sepals linear, lanceolate, flat, acuminate. Petals of
the same form, but twice as broad. Lip panduriform, slightly
hairy, with a cordate-ovate end, and near the base an elevated
ridge, which reaches half way down the lip and divides at the
point into two short plates. Column with a deep, purple. white-
edged, toothed hood, and short triangular wings.

“A rather pretty Brazilian Orchid, from Messrs. Loddiges.

30

We have only seen a single flower, beginning to wither. It was of
a yellow colour, not unlike in size and colour that of Miltonia
stellata, from which it differs in having an elevated plate passing
down the middle, narrower sepals, and a flat panduriform lip. It
is No. 1595 of Messrs. Loddiges’ Catalogue.”

Odontoglossum heterosepalum (Rchb.f.) Garay, stat nov.
Basionym: Odontoglossum angustatum var. heterosepa-
lum Rchb.f. in Linnaea 22: 850, 1850.

An examination of the holotype, FUNCK & SCHLIM 1242!
(W) collected in Merida convinces me that Reichenbach’s var-
iety constitutes a distinct species. It is hard to understand why
Reichenbach called this a variety of O. angustatum Lindl.
While his statement added to the description explains a famil-
iarity with the excessive natural variation observable within
species of Odontoglossum, the variation pattern of O. hetero-
sepalum cannot be accommodated within the pattern of varia-
tion of the southern O. angustatum.

3]

PLATE 1

Plate |. A. Psilochilus carinatus Garay. B. Pseudocentrum Purdii Garay

5

PLATE 2

Plate 2. A. Elleanthus Killipii Garay. B. Elleanthus formosus Garay.
32

PLATE 3

Plate 3. A. Cybebus grandis Garay. B. Sauroglossum distans Lindl. ex
Garay. C. Cyclopogon ovalifolium Pres|.

34

PLATE 4

Plate 4. A. Hapalorchis Lindleyanus Garay. B. Pelexia Hameri Garay.
C. Sarcoglottis Lehmannii Garay. D. Buchtienia rosea Garay.

35

PLATE 5

SS COD

Wwe

Plate 5. A. Platythelys peruviana Garay. B. Palmorchis colombiana Garay.

C. Eltroplectris pauciflora (Poepp. & Endl.) Garay.
36

PLATE 6

Plate 6. Epidendrum tropidioides Garay.
37

PLATE 7

Plate 7. A. Chondrorhyncha stenioides Garay. B. Osmoglossum pan-
duratum Garay.

38

BOTANICAL MUSEUM LEAFLETS VoL. 26, No. |
JANUARY 31, 1978

REVISION OF THE GENUS MASDEVALLIA - I.

HERMAN R. SWEET

The members of the genus Masdevallia have been in con-
stant demand during recent years. The genus is comprised of
some 400 species, most of which are poorly known, and those
in cultivation frequently misidentified. Several years ago I
started a complete revision of the genus, and this paper is the
first step towards its finalization.

In 1925 Kranzlin published a monograph of the genus Mas-
devallia which in reality was an updated version of THE GENUS
MASDEVALLIA by Miss Florence H. Woolward, originally pub-
lished in 1896. The best part of Kranzlin’s monograph is his
principal division of the genus into 14 sections. Here Kranzlin
demonstrated again, just as he had done with the genus On-
cidium, his ability to recognize natural groups. At the same
time he raised havoc with the assignment of individual species
into sections, a pattern quite uniform in all of his monographic
studies.

The first section taken up in this series 1s Triotosiphon pro-
posed by Schlechter in 1922, but overlooked by Kranzlin. It
was chosen purposely for it clearly reflects the taxonomic
status of the genus as a whole. Some of the species now
assigned to this section have been transferred to the genus
Physosiphon. Admittedly, the technical distinction between
Masdevallia and Physosiphon rests primarily in the calliferous
petals in Masdevallia; this particular character also dif-
ferentiates Masdevallia from the other genera of Pleurothal-
lidinae.

Because of the great similarity in gross morphology that
often exists among species belonging to the same section, it
became mandatory to dissect and draw every type specimen in
order to understand the true characters of their morphology.

When Schlechter chose the sectional name Triotosiphon —
translated as a tube having three ears — he emphasized the

39

nature of the flat but free sepaline lobes. Yet I believe that the
emphasis should be on the dorsally ventricose tube which at
once separates it from the Section Tubulosae.

The Section Triotosiphon contains eight species of which six
are new to science. Surprisingly I have seen no material from
Colombia or Peru, yet plants have been reported from Ven-
ezuela, Ecuador and Bolivia.

Section Triotosiphon (Schltr.) Sweet, comb. nov.
Basionym: Masdevallia subgen. Triotosiphon Schltr. in
Fedde, Rep. Beth. 10: 42, 1922.
Type: Masdevallia Bangii Schltr.

Small, epiphytic plants, densely caespitose. Roots filiform,
flexuous, glabrous. Secondary stem very short, completely
enclosed by tubular, imbricating sheaths. Leaves coriaceous,
very narrow. Inflorescence very slender, one-flowered. Flow-
ers rather small, yellow. Sepals connate into a dorsally ventri-
cose tube, suberect to arcuate, terminated by 3, free lobes;
lobes fleshy, more or less spreading, ear-like, hence the sec-
tional name, ecaudate. Petals, lip and column typical for the
genus.

KEY TO SPECIES

Ls. DAP IRtnCty DIODE 0460 brews een aae erage stake ade ev ers sees 2
Dae TA CRS se gia 5 hares Gy 298 220 eb a 9 bo HR eee oe Sa ea ee 4
2. Sepaline lobes ovate, obtuse; petals acute; lateral lobes of lip falcate-
Tia O lato s ps5 547 eee ease sone awe ean ee esse M. trioon
2a. Sepaline lobes lanceolate to narrowly ovate-lanceolate, acute; petals
ANCA, IR NIN ssc cao ores char den eee a RE ER ee ERA.
3. Lateral lobes of lip obliquely triangular, obtuse; terminal lobe of lip
ovate-rhombic; 3-dentate; disc glabrous ................. M. Bangii

3a. Lateral lobes of lip semi-elliptic; terminal lobe of lip cuneate-
subquadrate, erose-denticulate; disc at junction of terminal and lateral
lobes on each side with an obliquely inserted, semilunate callus .....
eee eee ee ee ee ee re wee eer ree er M. Lansbergii
4. Flowers arcuately curved; sepaline lobes triangular-lanceolate, sub-

acuminate; petals spathulate; lip with a pair of low ridges ......... |
4a. Flowers suberect; sepaline lobes ovate-lanceolate to linear-oblong,
acute to obtuse; petals linear-oblong; lip without ridges ........... 6

5. Sepaline lobes 8 mm. long; ridges of lip glabrous ... M. kyphonantha
Sa. Sepaline lobes 4 mm. long; ridges of lip pubescent . M. pseudominuta
6. Petals linear with a longitudinal, keel-like callus................-. 7

40)

6a. Petals dolabriform with globose calli; sepaline lobes 5 mm. long .....

Ca Sa Gie See Les CEA EEN S 5 CASO G MARE ORY REE nee OO aS M. gnoma
7. Sepaline lobes 5 mm. long; petals with a rounded, obscurely erose apex;
Le PACA ato teach des pen tea bores eee eesss M. irapana
7a. Sepaline lobes 10 mm. long; petals with a subtruncate apex, apiculate in
moiddles lip liNGSF cis seen ss cores a meri aware es cede M. venezuelana

Masdevallia Bangii Schltr. in Fedde, Rep. Beih. 10: 41, 1922.
Syn.: Physiphon Bangii (Schltr.) Garay in Can. Journ. Bot.
34: 249, 1956.
Type: Bolivia; La Paz; Coroico, Ungas, BANG 2424!
(AMES).

Epiphytic, caespitose plants, up to 5 cm. tall. Roots fascicu-
late, flexuous, glabrous. Secondary stem approximate, very
thin, up to 2 mm. long, one-leaved. Leaves fleshy, erect,
linear-oblanceolate, obtuse, gradually tapering toward the
base, up to 5 cm. long and 3 mm. wide. Inflorescence erect,
filiform, one-flowered, commonly shorter than the leaves, up
to 2cm. long. Bracts tubular, much shorter than the petiole, up
to 4mm. long. Flowers delicate, yellowish. Sepals connate into
a narrow, dorsally ventricose tube, up to 8 mm. in length, free
at apex; lobes lanceolate to narrowly ovate-lanceolate, dis-
tinctly carinate dorsally, acute, up to 3 mm. long and | mm.
wide. Petals obliquely linear-oblong, subtruncate at apex,
minutely apiculate in the center, very obscurely callose toward
base along anterior margin, up to 2.2 mm. long and 0.6 mm.
wide. Lip rather fleshy, from a cordate, oblong-cuneate base
3-lobed; lateral lobes obliquely triangular, obtuse; terminal
lobe ovate-rhombic, distinctly 3-dentate at apex, up to 2.2 mm.
long and | mm. wide. Column short with a three-lobed clinan-
drium, up to 2 mm. long. Ovary fusiform, articulate with the
pedicel, up to 10 mm. long.

Bolivia: La Paz; Coroico, Yungas, BANG 2424!(AMES, US).

Masdevallia gnoma, Sweet, sp. nov.
Type: Ecuador; Prov. Napo, vicinity of Papallacta, STACY
gs.n.! (AMES).

Epiphytica, minuta, usque ad 2.5 cm. alta; radicibus fas-
ciculatis, flexuosis, glabris; caulibus secundariis vix ullis, vag-

4]

inis 2, alte amplectentibus omnino obtectis; foliis carnosis,
lineari-oblanceolatis, acutis, basin versus sensim angustatis,
usque ad 2 cm. longis, 2 mm. latis; inflorescentia capillari,
uniflora, usque ad | cm. longa; floribus suberectis, parvis,
luteis vel luteo-viridibus; bracteis tubulosis, usque ad 2 mm.
longis; sepalis tubulosis, dorsaliter ventricosis, usque ad 5 mm.
longis, lobis liberis, ovato-lanceolatis, acutis, usque ad 4.5
mm. longis, | mm. latis; petalis inaequaliter dolabriformibus,
apice truncatis, in medio prominenter apiculatis, lamina antice
expansa, globoso-callosa, usque ad 2 mm. longis, 0.5 mm. latis;
labello oblongo-ligulato, antice subtruncato, margine erosulo;
disco lamellis triangulis ornato; toto labello usque ad 2.2 mm.
longo, 0.8 mm. lato; columna abbreviata, clavata, usque ad 2
mm. longa; ovario cylindrico, usque ad 2.5 mm. longo.

Epiphytic, minute plants, up to 2.5 cm. tall. Roots fascicu-
late, flexuous, glabrous. Secondary stem very short, com-
pletely enclosed by two loose-fitting, scarious sheaths, up to 2
mm. long. Leaves fleshy, linear-oblanceolate, acute, tapering
toward the base, up to 2 cm. long and 2 mm. wide. Inflores-
cence slender, one-flowered, up to | cm. long. Flowers sub-
erect, small, greenish yellow to yellow. Bracts tubulose, up to 2
mm. long. Sepals connate into a dorsally ventricose tube, 5
mm. long, free portions Ovate-lanceolate, acute, up to 4.5 mm.
long and | mm. wide. Petals unequally dolabriform, truncate
with a prominent, median apicule, the anteriorly expanded
blade witha globose callus, up to 2 mm. long and 0.5 mm. wide.
Lip oblong-ligulate, subtruncate in front with an erose margin;
disc with a pair of elevated triangular lamellae; whole lip up to
2.2 mm. long and 0.8 mm. wide. Column short, clavate, up to 2
mm. long. Ovary cylindric, up to 2.5 mm. long.

Ecuador: Prov. Napo; vicinity of Papallacta, STACY s.n.!
(AMES).

Masdevallia irapana Sweet, sp. nov.
Type: Venezuela; Sucre, Peninsula de Paria, N.E. of Irapa,
STEYERMARK 95079! (AMES).

Epiphytica, delicatula, caespitosa, usque ad 4 cm. alta;
radicibus fasciculatis, filiformibus, glabris; caulibus secun-
42

dariis abbreviatis, vaginis tubulosis, atro-brunneis omnino ob-
tectis, usque ad 5 mm. longis; foliis lineari-oblanceolatis, ob-
tusis, carnosis; basin versus sensim subpetiolatis, usque ad 5
cm. longis, 4mm. latis; pedunculo filiformi, unifloro, usque ad
15 mm. longo; bracteis tubulosis, | mm. longis; floribus
minutis, flavis vel aurantiacis; sepalis tubulosis, dorsaliter ven-
tricosis, usque ad 5 mm. longis; lobis liberis oblongo-ellipticis,
obtusis, usque ad 5 mm. longis, 1.4mm. latis; petalis lineari-ob-
longis, sub apice paulisper constrictis, apice rotundatis,
obscure erosis, margine anteriore callo lineari ornata, usque ad
2 mm. longis, 0.3 mm. latis; labello pandurato, dimidio basilar
carnoso, subquadrato, concavo, dimidio anteriori e cuneata
basi subrotondo, margine erosulo, toto labello usque ad 3 mm.
longo, 1.5 mm. lato; columna abbreviata, usque ad 2 mm.
longa; ovario cylindrico, usque ad | mm. longo.

Epiphytic, delicate, caespitose plants, up to 4cm. tall. Roots
fasciculate, filiform, glabrous. Secondary stem short, com-
pletely enclosed by imbricating, tubular, brown-purple
sheaths, up to 5 mm. long. Leaves linear-oblanceolate, obtuse,
tapering to a petiolate base, up to 5 cm. long and 4 mm. wide.
Peduncle very short, slender, up to 15 mm. long, one-flowered.
Bracts tubular, up to | mm. long. Flowers small, yellow to
orange in color. Sepals united into a dorsally ventricose tube,
up to 5 mm. long, free parts narrowly oblong-elliptic, obtuse,
up to 5 mm. long and 1.4 mm. wide. Petals linear-oblong,
somewhat constricted below the rounded, obscurely erose
apex, the anterior margin with a short, linear, longitudinal
thickening, up to 2 mm. long and 0.3 mm. wide. Lip pandurate;
basal part subquadrate thickened, concave; apical part from a
cuneate base subrotund with an erose margin, up to 3 mm. long
and 1.5 mm. wide. Column short, up to 2 mm. long. Ovary
cylindric, up to | mm. long.

Venezuela: Sucre, Peninsula de Paria, N.E. of Irapa,
STEYERMARK 95079! (AMES, VEN).

Masevallia Lansbergii Rchb.f. in Nederl. Kruidk. Arch. 4: 317,
1858.
Syn.: Physosiphon Lansbergii (Rchb.f.) L.O. Wms. in Bot.
Mus. Leafl. Harv. Univ. 7: 197, 1939.

43

Type: Venezuela; Caracas, LANSBERG 4! (W).

Epiphytic caespitose plants, up to 7 cm. tall. Roots fascicu-
late, glabrous. Secondary stem very short, completely en-
closed by two tubular sheaths, up to 5 mm. long. Leaves
linear-spathulate, obtuse, gradually tapering to base, subpetio-
late, up to 7 cm. long and 6 mm. wide, commonly smaller.
Peduncle filiform, one-flowered, up to 5 cm. long. Bracts tubu-
lar, up to 4mm. long. Flowers small, yellow. Sepals connate
into a dorsally ventricose tube, up to 8 mm. long, free part
linear-oblong, acute, up to 8 mm. long and 1.5 mm. wide. Petals
linear-oblong, truncate at apex with a small apiculum in mid-
dle, slightly tapering toward base, the anterior margin with a
small, subglobose callosity above the base, up to 2.5 mm. long
and 0.5 mm. wide. Lip 3-lobed, lateral lobes semi-elliptic;
terminal lobe cuneate-subquadrate, with an obtuse, subtrun-
cate, erose-denticulate apex, at the junction of the terminal
lobe and lateral lobes on each side provided with an obliquely
inserted, semi-lunate callus; whole lip up to 3 mm. long and |
mm. wide. Column short with a distinct foot, cylindric, up to 2
mm. long. Ovary cylindric, up to 2 mm. long.

Venezuela: Caracas, LANSBERG 4! (W); Tovar, FENDLER
1369! (AMES)

Masdevallia kyphonantha, Sweet, sp. nov.
Type: Venezuela; Estado Yaracuy; 7 km. N. of Salom,
STEYERMARK 106263! (AMES).

Epiphytica, caespitosa, usque ad 5 cm. alta; radicibus fasci-
culatis, flexuosis, glabris; caulibus secundariis approximatis,
valde abbreviatis, vaginis 2, ample tubulosis omnino obtectis,
usque ad 4 mm. longis; foliis carnosis, lineari-oblongis, paulis-
per basin versus attenuatis, obtusis, usque ad 5 cm. longis, 2
mm. latis; inflorescentia capillari, uniflora, usque ad 2.5 cm.
longa; bracteis tubulosis, usque ad 4 mm. longis; floribus al-
bidis; sepalis tubulosis, dorsaliter ventricosis, arcuatis, inde
nomen, usque ad 18 mm. longis, lobis liberis, lanceolato-trian-
gularibus, subacuminatis, usque ad 8 mm. longis, 3 mm. latis;
petalis oblique spathulatis, apice rotundatis, minute muc-
ronatis, Margine anteriore in medio callo globoso ornata, usque

44

ad 3.5 mm. longis, 1.2 mm. latis; labello oblongo-ligulato,
antice rotundato vel subtruncato, erosulo, disco longitudinali-
ter bicarinato, usque ad 5 mm. longo, 2 mm. lato; columna
paululo arcuata, subclavata, usque ad 3.5 mm. longa; ovario
cylindrico, usque ad 5 mm. longo.

Epiphytic, caespitose plants, up to 5 cm. tall. Roots fascicu-
late, flexuous, glabrous. Secondary stem approximate, very
short, completely enclosed by two, loose-fitting, tubular
sheaths, up to 4 mm. long. Leaves fleshy, linear-oblong,
somewhat tapering toward the base, obtuse, up to 5 cm. long
and 2 mm. wide. Inflorescence very slender, one-flowered, up
to 2.5 cm. long. Bracts tubular, as long as the pedicels, up to 4
mm. long. Flowers whitish. Sepals connate into a dorsally
ventricose, arcuately curved tube, hence the name, up to 18
mm. long, the free portions lanceolate-triangular, subacumi-
nate, up to 8 mm. long and 3 mm. wide. Petals obliquely
spathulate, rounded at apex, minutely mucronate, with an
obscurely globose callus near the middle of the anterior mar-
gin, up to 3.5 mm. long and 1.2 mm. wide. Lip oblong-ligulate,
rounded to subtruncate in front with an erose margin; disc
provided with a pair of elevated, longitudinal ridges, up to 5
mm. long and 2 mm. wide. Column slightly arcuate, subcla-
vate, up to 3.5 mm. long. Ovary cylindric, up to 5 mm. long.

Venezuela: Estado Yaracuy; 7 km. N. of Salom, STEYER-
MARK 106263! (AMES, VEN).

Masdevallia pseudominuta, Sweet, sp. nov.
Type: Venezuela; Curimagua, Sierra de San Luis, GARCIA
s.n. sub DUNSTERVILLE 538! (AMES).

Epiphytica, caespitosa, usque ad 3 cm. alta; radicibus fili-
formibus, flexuosis, glabris; caulibus secundartis valde abbre-
viatis, seu vix ullis, vaginis imbricantibus, tubulosis obtectis;
foliis carnosis, lineari-oblanceolatis, usque ad 3 cm. longis, 2
mm. latis; pedunculo gracili, unifloro, usque ad 2 cm. longo;
bracteis tubulosis, usque ad 3.5 mm. longis; floribus luteis;
sepalis tubulosis, dorsaliter ventricosis, arcuatis, 6mm. longis,
lobis liberis, triangulari-lanceolatis, subacuminatis, usque ad 5
mm. longis, | mm. latis; petalis oblique spathulatis, obtusis,

45

obscure apiculatis, margine anteriore in medio obscure sub-
globosa, usque ad 2 mm. longis, 0.8 mm. latis; labello lineari-
oblongo, subtruncato, disco longitudinaliter bicarinato, carinis
puberulentibus, usque ad 2.5 mm. longo, 0.8 mm. lato; colum-
na abbreviata, clavata, usque ad 2 mm. longa; ovario cylindri-
co, usque ad 2 mm. longo.

Epiphytic, small, caespitose plants, up to 3 cm. tall. Roots
filiform, flexuous, glabrous. Secondary stem short, completely
enclosed by imbricating, tubular sheaths. Leaves fleshy, lin-
ear-oblanceolate, up to 3. cm. long and 2 mm. wide. Peduncle
slender, one-flowered, up to 2 cm. long. Bracts tubular, up to
3.5mm. long. Flowers yellow. Sepals are united into a dorsally
ventricose, somewhat arcuately curved tube 6 mm. in length,
free portions spreading, triangular-lanceolate, subacuminate.,
up to5 mm. long and | mm. wide. Petals obliquely spathulate,
obtuse, obscurely apiculate with an obscurely subglobose cal-
lus near the middle of the anterior lobe, up to 2 mm. long and
0.8 mm. wide. Lip linear-oblong, subtruncate in front with a
pair of longitudinal, puberulent ridges, up to 2.5 mm. long and
0.8 mm. wide. Column short, clavate, up to 2 mm. long. Ovary
cylindric, up to 2 mm. long.

Venezuela: Curimagua, Sierra de San Luis, GARCIA s.n. sub
DUNSTERVILLE 538! (AMES).

Masdevallia trioon Sweet, sp. nov.
Type: Ecuador; Condor, FISKE s.n.! (AMES)

Epiphytica, perpusilla, caespitosa, usque ad 4 cm. alta;
radicibus filiformibus, flexuosis, glabris; caulibus secundariis
vix ullis, vaginis tubulosis, 2, omnino obtectis, usque ad 2 mm.
longis; foliis lineari-spathulatis, obtusis, basin versus angus-
tatis, usque ad 4 cm. longis, 3 mm. latis; pedunculo filiform,
unifloro, usque ad 1.5 cm. longo; bracteis tubulosis, usque ad 3
mm. longis; floribus luteis; sepalis dorsaliter ventricosis, usque
ad 6 mm. longis, lobis liberis patentibus, ovatis, obtusis, inde
nomen, usque ad 2 mm. longis, 1.6 mm. latis; petalis e cuneata
basi oblongo-ellipticis, acutis, margine anteriori callo globoso
decora, usque ad 3 mm. longis, | mm. latis; labello e cuneato-
subquadrata basi 3-lobo, lobis lateralibus falcato-triangulari-

46

bus, lobo terminali rectangulariter-oblongo, recurvo, margine
integro, toto labello, usque ad 3 mm. longo, | mm. lato; col-
umna erecta, usque ad 3.5 mm. longa; ovario cylindrico, usque
ad 3.5 mm. longo.

Epiphytic, small, caespitose plants, up to 4 cm. tall. Roots
filiform, flexuous, glabrous. Secondary stem hardly any, com-
pletely enclosed by two tubular sheaths, up to 2 mm. long.
Leaves linear-spathulate, obtuse, tapering to base, up to 4cm.
long and 3 mm. wide. Peduncle filiform, one-flowered, up to
1.5cm. long. Bracts tubular, up to 3 mm. long. Flowers yellow.
Sepals united into a dorsally ventricose, inflated tube, up to 6
mm. long, free lobes spreading, ovate, obtuse, up to 2 mm. long
and |.6 mm. wide. Petals from a cuneate base oblong-elliptic,
acute, along anterior margin below middle provided with a
small, globose callus, up to 3 mm. long and | mm. wide. Lip
from a cuneate-subquadrate base 3-lobed; lateral lobes fal-
cate-triangular; midlobe rectangular-oblong, recurved with en-
tire margin; whole lip up to 3 mm. long and | mm. wide.
Column erect with a short foot, up to 3.5 mm. long. Ovary
cylindric, up to 3.5 mm. long.

Ecuador: Condor, FISKE s.n.! (AMES).

Masdevallia venezuelana Sweet, sp. nov.
Type: Venezuela; Rancho Grande, PITTIER 13966!
(AMES).

Epiphytica, caespitosa, usque ad 10 cm. alta; radicibus fas-
ciculatis, filiformibus, flexuosis, glabris; caulibus secundariis
abbreviatis, vaginis tubulosis omnino obtectis, usque ad 6 mm.
longis; foliis carnosis, oblanceolato-spathulatis, obtusis, basin
versus in petiolo angustatis, usque ad 6 cm. longis, 8 mm. latis,
vulgo minoribus; pedunculo gracili, unifloro, usque ad 8 cm.
longo; bracteis tubulosis, pedicellis aequilongis, usque ad 5
mm. longis; floribus citrinis vel aurantiacis, satis magnis;
sepalis dorsaliter ventricosis, 10 mm. longis, lobis liberis linea-
ri-oblongis, obtusis vel rotundatis, usque ad 10 mm. longis, 2
mm. latis; petalis hyalinis, linearibus, apicem versus paululo
dilatatis, subtruncatis, in medio apiculatis, margine anteriori
callo cariniformi ornata, usque ad 3 mm. longis, 0.5 mm. latis;

47

labello luteo, lineari, parte basali carnosa, excavata, in medio
marginis in disco decurrentibus, parte apicali membranacea,
oblongo-lineari, apice erosa, toto labello usque ad 3.5 mm.
longo, | mm. lato; columna erecta, cylindrica, usque ad 2 mm.
longa, ovario cylindrico, sulcato, usque ad 2 mm. longo.

Epiphytic, caespitose plants, up to 10cm. tall. Roots fascicu-
late, filiform, glabrous. Secondary stem short, completely en-
closed by tubular sheaths, up to 6 mm. long. Leaves fleshy,
oblanceolate-spathulate, obtuse, tapering to a distinct petiole,
up to6cm. long and 8mm. wide, commonly smaller. Peduncle
slender, one-flowered, up to 8cm. long. Bracts tubular, as long
as the pedicel, up to 5 mm. long. Flowers lemon-yellow to
golden yellow, large for the section. Sepals connate into a
dorsally ventricose tube, 10 mm. in length, free parts linear-
oblong, obtuse to rounded at apex, up to 10 mm. long and 2
mm. wide. Petals hyaline, linear, somewhat expanded at apex,
subtruncate, apiculate in middle, the anterior margin with a
long, fleshy keel, up to 3 mm. long and 0.5 mm. wide. Lip
yellow, linear, basal half fleshy, excavate, with the margins
decurved in middle onto the disc; apical half membranaceous,
oblong-linear with erose apex, up to 3.5 mm. long and | mm.
wide. Column erect, cylindric, with a distinct foot, up to2 mm.
long. Ovary cylindric, sulcate, up to 2 mm. long.

Venezuela: Rancho Grande PITTIER 13966! (AMES, US),
DUNSTERVILLE 528! (AMES), STEYERMARK 89793! (AMES,
VEN), PITTIER 11820! (AMES, G, US).

48

PLATE 8

Plate 8. A. Masdevallia gnoma Sweet. B. Masdevallia kyphonantha Sweet.
C. Masdevallia pseudominuta Sweet.

49

PLATE 9

Apex ef Leof ‘ :

Section trough Leaf

Perot & Lip

(Lip spread)

eM

Column & lip

Plate 9. Masdevallia irapana Sweet.

nm
oy
4

PLATE 10

Plate 10. Masdevallia gnoma Sweet. B. Masdevallia Bangii Schltr.,
C. Masdevallia trioon Sweet.

8

PLATE 11

ame

a_i

Column Foce

ae es
Una de =} Mure ”
inderside Roste An ther & Peilinia

saa NY

Column & Lip

Plate 11. Masdevallia Lansbergii Rchb.f.

52

PLATE 12

ee i...
n mM,
eral | Column face dlaaf
: ' : be
an if BR aN ail |i
Vt '
1 ’ aa
ey

Upper & Mid- leaf

Plate 12. A. Masdevallia kyphonantha Sweet. B. Masdevallia pseudominu-
ta Sweet.

bh

PLATE 13

Vener

Lip

a orem faecteses, 7,
! ee
fom Petal & Lip

0

Anther & Pollina

Som

Plate 13. Masdevallia venezuelana Sweet.

54

BOTANICAL MUSEUM LEAFLETS VoL. 26, No. |
JANUARY 31, 1978

STUDIES IN THE GENUS ONCIDIUM — II
SECTION SERPENTIA

JOHN E. STACY

As I have stated in the first paper of this series, the various
sections of Oncidium will be handled in a random sequence,
mostly dictated by existing needs for clarification of cultivated
material.

The Section Serpentia was established by Garay in 1970 in
his review of the genus Oncidium. The group of plants compris-
ing this section had already been brought together by Kranzlin
in 1922 as a subsection notwithstanding the lack of any phyletic
or genetic proximity with the rest of the Section Varicosa to
which it was attached.

The outstanding characteristic of this section is the remarka-
ble habit of the plants. The inflorescences which may be com-
pared with those of O. volvox, O. acinaceum, O. Schillera-
num to mention a few, often develop adventitious plantlets in
lieu of flowers. The result is a twining habit which, without
closer scrutiny, appears to have plants situated distantly on
thin rhizomes or stolons. The true nature of these plants was
first recognized by Dr. Rodrigues in Costa Rica, who together
with Dr. Daniels, give an excellent discussion on the morpho-
logy of these plants in ORQUIDEOLOGIA 7: 79-84, 1972.

Although the Section Serpentia contains only a handful of
species, the individual members are still poorly known, most
probably due to the great similarity in their vegetative growth.
The type-species, O. globuliferum, for a long time remained
unknown, but it is available for study now in the Paris her-
barium. When Reichenbach first examined Humboldt’s mate-
rial he hastily united his own O. Scansor with it. Yet both
plants are amply distinct from one another as will be shown
later. To make matters worse, Reichenbach described a new
variety of O. globuliferum from Costa Rica which was indeed
different from his concept of O. Scansor.

55

Perhaps the greatest mystery in this group of plants is O.
serpens Lindl. Lindley based his description on a single dried
specimen without actually examining the lip for details. Since
the type-specimen had only a single flower very nicely pressed,
an examination of it without boiling it first, gave rise to a wrong
description of its shape and the nature of its callus. Through the
courtesy of Dr. Peter Taylor of the Orchid Herbarium, Royal
Botanic Gardens, Kew, I have studied the type-specimen and |
find it identical with O. trachycaulon Schltr. and O. Sancti-
Pauli Kral.

While the plants of the species mentioned above all have
smooth peduncles without nodes which twine about branches,
Oncidium Harlingii, a newly discovered species, exhibits a
very different habit. The plants are epiphytic on Selaginella
growing on the ground, its peduncles are upright and jointed.
The species is endemic in Ecuador and its distribution is limited
in the area of Gualaceo to General Plaza where it has been
collected on three different occasions in association with
Odontoglossum crinitum.

Section Serpentia (Krzl.) Garay in Taxon 19: 455, 1970.
Type: Oncidium serpens Lindl.

The flowers are medium to large for the plant and range from
2 to5cm. vertically, yellow in color with reddish brown macu-
lations on the sepals, petals and base of lip. The sepals and
petals are clawed, spreading; the petals somewhat wider than
the sepals. The lip is most conspicuous, usually with small
basal lobes; the midlobe is emarginate, transversely reniform,
solid-colored except at the base, with a variously tuberculate
crest. The column is short, auriculate, and thickened below the
stigma; the rostellum is short.

The plants are small, epiphytic, with remotely-spaced
pseudobulbs on slender, wiry, twining or erect inflorescences.
The mature pseudobulbs are monophyllous, rarely longer than
3 cm., round to ovate, slightly compressed, enclosed by dis-
tichously imbricating, leafy sheaths. The secondary inflores-
cence is short, emerging from the axils of the sheaths, fre-
quently from less developed pseudobulbs, usually single-

56

flowered or rarely few-flowered on a fractiflex rachis. The
leaves are oblong-ovate, acute or rounded at apex and witha
conduplicate petiole.

KEY TO SPECIES

Inflorescence many-jointed, covered with adpressed, chartaceous

sheaths; petals larger than sepals .................... QO. Harlingii
. Inflorescence without joints or sheaths; petals more or less similar to
BEERS. ikea aes 6 owt AS ees eee eee bee eee ewe Bes 2
Leaves rounded at apex; column wings quadrate, entire; clinandrium
OTE VOU esate Parnas Ca dd hs CERO VA Ee EEA OEE A 3
. Leaves pointed at apex; column wings serrate; clinandrium strongly
i NE oc se ee oe ee oa Boe eens aes eee ey oun 4 O. serpens
Midlobe of lip sessile, overlapping lateral sepals ........ O. Scansor
. Midlobe of lip produced from an elongate isthmus, exposing lateral

SOUS 4 ccc eaey Eis hae GORY Ree cas be Nae ee eae O. globuliferum

SYSTEMATICS OF SPECIES

Oncidium globuliferum H.B.K., Nov. Gen. et Sp. Pl. 1: 347,
1816.

Type: Colombia; Dept. Cauca, near Lake Timbio, Hum-
boldt 2072! (P).

Syn.: Oncidium convolvulaceum Lindl. in Lindl. & Paxt.,
Fl. Gard. 1: 102, 1850.

Type: Venezuela; without precise locality, Funck & Schlim
1444! (K-L).
Oncidium globuliferum var. costaricense Rchb.f. in
Gard. Chron. 1678, 1871.

Type: Costa Rica; without precise locality, Zahn 149! (W).
Oncidium Wercklei Schltr. in Fedde, Rep. Beih. 19:
68, 1923.

Type: Costa Rica; San Jeronimo, Werckle 46! (AMES).

Observation: Sepals up to 15 mm. long, 5 mm. wide, obo-

vate-oblong, slightly clawed at the base; lateral sepals free and
slightly narrower. Petals subsimilar, up to 15 mm. long and 6
mm. wide. Lip is conspicuous, 3-lobed; basal lobes small,

57

suborbicular; midlobe from a prominent isthmus transversely
reniform, emarginate, up to 3.5 cm. wide and 2.8 cm. long
including the base. Pseudobulbs somewhat compressed, round
to ovate, spaced remotely on slender, wiry growths. Peduncle
either lateral from base of pseudobulb or from developing
plantlet, usually single-flowered but specimens have been seen
with several-flowered, fractiflex rachis; the largest pseudobulb
up to 3.5 cm. long with a leaf up to 10 cm. long.

DISTRIBUTION: Costa Rica south to Bolivia.

Oncidium Harlingii Stacy, sp. nov.
Type: Ecuador; Gualaceo 30 K. West of Limon, growing on
ground among moss at 2300 m., HARLING 12835!
(GB).

Epiphytica, erecta, simplici vel ramosa; pedunculis,
teretibus, satis rigidis, multi-articulatis, vaginis ad-pressis, jam
emarcidis, internodiis sesquipollicaris subaequilongis vestiti;
pseudobulbis valde distantibus, leviter compressis, ovoideo-
lenticularibus, vaginis distichis, foliiferis omnino obtectis, un-
ifoliatis, usque ad 2 cm. altis; foliis in exsiccatione satis
papyracels, Ovato-lanceolatis, acutis vel subacuminatis, basin
plus-minusuve petiolatis, petiolo usque ad 1.5 cm. longo in-
cluso usque ad II cm. longis, 2.5 cm. latis; inflorescentiis
abbreviatis, 1-3 floris; bracteis cucullato-lanceolatis,
acuminatis, Ovariis pedicellatis brevioribus; floribus con-
spicuis; sepalo postico obovato-elliptico, obtuso, paulo con-
vexo, usque ad 8 mm. longo, 5 mm. lato; sepalis lateralibus
inter se breviter connatis, patulis, ovato-oblongis, acutis,
usque ad 10 mm. longis, 4mm. latis; petalis quam sepalis multo
majoribus, ovato-ellipticis obtusis vel rotundatis, usque ad 12
mm. longis, 7 mm. latis; labello sessili, basin utrinque au-
riculato, subrotundo, deinde in laminam transverse renifor-
mem, antice emarginatam seu bilobam expanso; disco basin
callo triquetro, tota superficie minutissime muricato ornato;
toto labello 26 mm. longo, 30 mm. lato, basin 11 mm. lato;
columna humill, juxta stigma auriculata, auriculis oblique sub-
quadratis; ovario pedicellato cylindrico, usque ad 2 cm. longo.

Observation: O. Harlingii is readily separated from the other
58

species belonging to this section by the rather straight and erect
growths with adpressed sheaths at the many nodes and by the
triquetrous callus at the base of the lip. The dorsal sepal is
obovate, obtuse, up to 8 mm. long and 5 mm. wide; the lateral
sepals are shortly connate, spreading, ovate-oblong, up to 10
mm. long and 4 mm. wide. The petals are ovate elliptic, obtuse
or rounded, up to 12 mm. long and 7 mm. wide. The lip ts
sessile, auriculate at the base with subrotund lobes, up to I1
mm. across; the midlobe is transverse-reniform, emarginate at
the apex, up to 2.6 cm. long and 3 cm. wide; the callus is small,
triquetrous, with minute superficial processes along the mar-
gins. The column is short with obliquely subquadrate auricles
on either side of the stigma.

DISTRIBUTION: Endemic in southern Ecuador.

Oncidium Scansor Rchb.f. in Linnaea 22: 844, 1850.
Type: Venezuela; Prov. Merida at 2000 m., Funck & Schlim
1022! (W).

Observation: O. Scansor is readily separated from O.
globuliferum by the midlobe of lip being sessile; the lateral
lobes are small and together form a somewhat saggitate base.
The lip is up to 3.8 cm. long and 5 cm. wide, up to 15 mm. across
the lateral lobes; the flat basal portion of the callus is pubes-
cent. The dorsal sepal is elliptic-oblong, acute, up to 16 mm.
long and 6 mm. wide; the lateral sepals are similar and slightly
oblique, up to 17 mm. long and 5 mm. wide. The petals are
obovate, up to 18 mm. long and 7 mm. wide. The column wings
are subquadrate.

DISTRIBUTION: Venezuela, Colombia.

ILLUSTRATION: Dusnterville & Garay, Venez. Orch. Ill. 2:
249, 1961 as O. globuliferum.

Oncidium serpens Lindl., Gen. and Sp. Orch. Pl. 204, 1833.
Type: Ecuador; Prov. Esmeraldas, Patacocha, at 1900 m.,
Hall 4! (K).
Syn.: Oncidium trachycaulon Schltr. in Fedde, Rep. Beih.
7: 191, 1920.

59

Type: Colombia; Dept. Cauca, Madero, s.n.

Observation: The plants of this species are readily separated
from O. Harlingii by the absence of sheaths and nodes along
the growths. The lip of the flower is longer than wide, and in the
type-flower it is more or less oblong-pandurate with an auricu-
late base. It measures up to 12 mm. long and 8 mm. across the
midlobe, but may be somewhat wider. The callus, as recon-
structed from the type-specimen, is tuberculate and it is dif-
ficult to explain Lindley’s interpretation of it being bilamellate.
The sepals and petals are clawed, elliptic-obovate, up to 11
mm. long and 5 mm. wide; the petals are less prominently
clawed. The column is up to 5 mm. long with small, quadrate,
serrated wings; the clinandrium is conspicuously cucullate,
truncate. The pedicellate ovary is up to 18 mm. long and the
floral bracts are up to 4mm. long. The leaves are ovate, acute,
up to S cm. long and 1.5 cm. wide. The somewhat compressed
pseudobulbs are oblong-ovate and rarely more than 2.5 cm.
long. The inflorescence is usually several-flowered.

DISTRIBUTION: Colombia. Ecuador.

ACKNOWLEDGMENTS

To Dr. Peter Taylor, the Orchid Herbarium, Royal Botanic
Gardens, Kew, England, I am indebted for permission to exam-
ine type-material. I wish to express my appreciation also to Dr.
R. Lucas Rodriguez of the University of Costa Rica for his
constructive comments on the morphology of species referable
to this Section, to Dr. Leslie A. Garay of the Orchid Herbarium
of Oakes Ames for his continuing guidance, and to Miss Ger-
trude Ahern for her secretarial assistance.

60

PLATE 14

Plate 14. Oncidium Scansor Rchb.f.

61

PLATE 15

CN at

Plate 15. Oncidium Harlingii Stacy

62

PLATE 16

Plate 16. A. Oncidium Harlingii Stacy. B. Oncidium serpens Lindl. C. On-
cidium globuliferum H.B.K.

63

BOTANICAL MUSEUM LEAFLETS

HARVARD UNIVERSITY

1978 VoL. 26, No. 2-4

BIBLIOGRAPHY OF PRECAMBRIAN
PALEONTOLOGY AND PALEOBIOLOGY

*STANLEY M. AWRAMIK AND **ELso S. BARGHOORN

This bibliography was started in 1968 while the first author
was a graduate student at Harvard University, and has been
carried on as a continuing project since that time, primarily at
the University of California, Santa Barbara. Since the litera-
ture in this relatively new, but rapidly expanding area of
paleontology is growing at an increasing rate, it was decided to
make the bibliography available at this time to all workers in
this and related fields.

In selecting the references, an effort was made to exclude
material which is purely, or largely, geochemical in context,
and to avoid overlap with the comprehensive bibliography on
‘*Chemical Evolution and the Origin of Life’’. The latter has
been published in Space Life Sciences 2 (1970): 225-295; 3
(1973): 293-304; 4 (1973): 309-328; and in Origins of Life 5
(1974): 507-527; 6 (1975): 285-300; 7 (1976): 75-85; 8 (1977):
59-65.

An effort has been made also to avoid duplication insofar as
possible, of references which are indexed in the volume
Stromatolites (edited by M.R. Walter, Elsevier, 1976), which
contains a comprehensive bibliography of published work on
stromatolites, both Precambrian and Phanerozoic.

* Dept. of Geological Sciences, University of California, Santa Barbara, California,

93100.
** Department of Biology, Harvard University, Cambridge, Mass. 02138

Published monthly except during July and August by the Botanical Museum, Harvard University,
Cambridge, Massachusetts 02138. Printed by Harvard University Printing Office. Subscription:
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1974 — Stratigraphy of the upper Proterozoic on the northwestern slope of
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1964 Columnar stromatolites from the Riphean to north of the Siberian
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1969 Stromatolity verkhnego dokembriya severa sibirskoi platformy 1
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1964 —_ Division of the Riphean based on stromatolites. In Precambrian
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1965 Procedure for studying Conophyton stromatolites and their strat ig-

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1968 Detailed stratigraphy of the Upper Proterozoic based on stromato-
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Komar, V.A., M.A. Semikhatoy, and S.N. Serebryakov

1973 Characteristic range of stromatolite species from the Riphean
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Komar, V.A., M.A. Semikhatov, S.N. Serebryakov, and B.G. Voronoy

1970 Novie dantie po stratigrafii i istorii razvitiya rifeya yugo-
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Komar, V.A. and S.N. Serebryakoy

1969 K stratigrafii dokembriya Turykhanskogo podnyatiya. Dokl.

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1974 Normal alkanes in petroleum of Vendian and Cambrian deposits of

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1972 Some new microorganisms from the Bohemian Precambrian (Up-
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1973 Algal colony and rests of other microorganisms in the Bohemian
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1974 Some new results of the Precambrian research in Czechoslovakia.
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1974 Acritarchs from the Bohemian Precambrian (Upper Proterozoic)
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Kopeliovich, A.V. and I.N. Krylov.

1960 =O strukturakh rastvoreniya v stromatolitakh. (On solution struc-

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1969 The geological history of the earliest algae and their stratigraphical
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Korde, K.B.

1950 Mikroskopicheskaya struktura nasloenii stromatolitov 1 tipy sok-
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1953 Formation and systematic position of the conical and cylindrical
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1953 Concerning certain questions on the study of fossil algae (in Rus-
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1958 Concerning several species of fossil blue-green algae (in Russian).
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Korolev, V.G. and I.N. Krylov

1962 Contribution to the stratigraphy of the Upper Precambrian of the
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Korolyuk, I.K.

1957. Certains Stromatolites du Cambrien de amphitheatre d’Irkoutsk.
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1958 = Znachenie stromatolitoz dlya stratigrafil kKembriya i dokembrya na
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1960 = Stromatolity nizhnego kembriya i proterozoya Irkutskogo am-
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1960 = Subdivision of the Cambrian and Precambrian in eastern Siberia
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1960 Konofitony v uluntuiskoi svite Pribaikalya. Materialy po geol. i
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1963 Stromatolites of the Upper Precambrian. In Upper Precambrian
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1966 Mikroproblematika rifeya i nizhnego kembriya Pribaikalya 1

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1968 Biogermiye obrazovan iya zapadnogo Pribaikalya (in Russian).
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1962. Znachenie stromatolitov dlya stratigrafii rifeya. In Soveshehanie
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Korolyuk, I.K. and I.A. Korillova

1973 Lithology of limestone bioherms of the lower Permian Shakhtau
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Korolyuk, I.K. and N.S. Lagutenkova

1965 Further finds of problematic microfossils in Bavly sediments.

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1965 — Stromatolity i Mikroproblematika Pribaikalya. In Vsesoyuzn. sim-
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1969 Stromatolity motskoi svity yuzhnogo pribaikal’ya 1 yugo-vosto-
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1971 Stromatolity Yuzhnogo Pribaikalya (in Russian). (Stromatolites of
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Koshchevoi, V.V.

1973 Nekotorye novye iskopaemye mikroorganizmy dokembriya (in
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1975 Nekotorye momenty izucheniya mikroorganizmov dokembriya s
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Koslov, V.

1973 The structure of the Upper Proterozoic Ukskian and Ashinian
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Krasilshehikov, A.A., N.P. Golovanov, and V.E. Milshtein

1965 Kstratigrafii verkhneproterozoiskikh otlozhenii raiona
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1936 Drevneishie vodorosli Polshe-Pitskogo raiona Eniseiskei Taigi.

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1937. Vodorosli i arkheotsiaty drevneishikh tolsheh Potekhinskogo
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1945 Paleontological data on the correlation of some pre-Cambrian se-
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1946 Nekotorye stromatolity proterozoya Kabyrzinskogo raiona Gor-
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1956 Vodorosli. In Atlas rukovodyashehikh form iskopaemoi fauny i
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1924. Archaeoxylon Krasseri, ein Pflanzenrest aus dem bohmischen

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1965 Stromatolite von Tsumeb, Siidwest Afrika. Sci. Soc. Journ., pp.

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1965 | Mikrospory v ostatochnykh zheleznykh rudakh Krivorozhskogo 1
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Krendelov, R. and A. Luchko

1970 Medistye konglomeraty i peschaniki kyllaskogo khrebta (Y ugo-
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Krishna Murty, M.

1972 Stromatolites from the Bijawars of the Joga Area, Hoshangabad

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1969 Stromatolites and oncolites in the Otavi series, South West Africa.

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1959 Stromatolites from the Riphean of the Urals (in Russian). Do&l.
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1959 Rifeyskiye Stromatolity Ostova Kil’dina. (Riphean stromatolites
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1960 printsipakh sistematiki rifeiskikh stromatolitov. Byull. Mosk.
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1960 Oznacheniistromatolitov Collenia buriatica Mas\. dlya stratigrafil
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1960 O razvitii stolbehatykh vetvyashehikhsya stromatolitov v rifee
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|,

1963

1964

1964

1965

1966

1966

1966

1967

1967

1968

1968

1969

1921

1972

1972

Ramifying columnar stromatolites of the Riphean of the Southern
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Pervyi Vsesoyuznyi kollokvium po stromatolitam onkolitam i
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O zakonomernostyakh izmenchivosti morfologicheskikh priz-
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Znachenie stromatolitov dlya stratigrafii dokembriya (in Russian).
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Rifeiskiye i nizhnekembriiskiye stromatolity Tyan-Shanya i
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Stromatolity pogranichnykh sloev dokembriya i nizhnego kem-
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Znachenie stromatolitov dlay Problemy granitsy Dokembriya i
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Sravnitelnaya kharakteristika stromatolitov dokembriya i kem-
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Znachenie stromatolitov dla stratigrafii verkhnego dokembriya
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1974 Natural evidence for chemical and early biological evolution. Ori-
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Langille, G.B.

1974 Earliest Cambrian: Latest Proterozoic ichnofossils and problema-
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175

BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY

MASSACHUSETTS, May 30, 1978 — VOL. 26, No. 5

CAMBRIDGE,

De Plantis Toxicariis e Mundo
Novo Tropicale Commentationes XXIII

NOTES ON BIODYNAMIC PLANTS OF ABORIGINAL USE
IN THE NORTHWESTERN AMAZONIA

RICHARD EVANS SCHULTES

This article continues a series of notes on plants employed in
the rich ethnopharmacopeas of the Indians of the Amazon
Valley — mainly in Colombian, Ecuadorian and Peruvian ter-
ritories. The primary purpose of this series is manifestly to call
to the attention of phytochemical and pharmacological re-
search specialists the hitherto neglected but extensive knowl-
edge of biodynamic plants amongst the aboriginal populations
of these regions — a knowledge destined to disappear rapidly in
the face of threatening or even presently active acculturation.

Some of the notes in the following pages have come from my
own ethnobotanical studies in the northwest Amazon from
1941 to the present. I have, however, drawn also upon the field
research of colleagues, especially upon the studies of my
former student, Dr. Homer V. Pinkley, amongst the Kofan
Indians of Ecuador and Colombia and upon the rich store of
knowledge of my Colombian colleague, Prof. Hernando
Garcia-Barriga of the Instituto de Ciencias Naturales, Univer-
sidad Nacional, Bogota, Colombia. I have further incorporated
into these notes several of the preliminary results of the Al-
pha-Helix Amazon Expedition 1976-77, Phase VII, dedicated
to Ethnopharmacological Studies of the Flora and Fauna of the

Published monthly except during July and August by the Botanical Museum, Harvard University,
Cambridge, Massachusetts 02138. Printed by Harvard University Printing Office. Subscription:
$25.00 a year, net, postpaid. Orders should be directed to Secretary of Publications at the above
address. Second-Class Postage Paid at Boston, Massachusetts.

Pebas Region of the Peruvian Amazon, and I have freely
quoted from the final report (unpublished) of this expedition.

I am pleased to acknowledge the collaboration of my Colom-
bian colleagues, Prof. Alvaro Fernandez-Perez and Dr.
Enrique Forero of the Herbario Nacional de Colombia, and of
Prof. Oscar Tovar of the Universidad Nacional Mayor de San
Marcos, Lima, as well as the specialists who have identified
many of the voucher specimens.

The notes in the following pages are arranged in accordance
with the Engler & Prantl system. Voucher specimens cited are
deposited in the Economic Herbarium of Oakes Ames, Botani-
cal Museum of Harvard University, the Gray Herbarium of
Harvard University, the Herbario Nacional de Colombia in
Bogota, the Herbario del Museo de Historia Natural, Lima or
in several of these institutions.

It is appropriate to make acknowledgment of the role of the
National Science Foundation which, through Grant No.
DEB75-20107, has made possible some of the research re-
ported in the following pages. The work on the Alpha-Helix
Expedition was possible through Grant No. DEB72-02536-
Garey 6-446406-21412.

HAEMODORACEAE

Schiekia orinocensis (HBK.) Meissner, Pl. Vasc. Gen. Comm.
(1842) 300.

CoLomBiaA: Comisaria del Vaupes, Rio Vaupes, Cerro de Mitu. *‘On
granite rocks. Flowers deep yellow.’*> September 7, 1951. R.E. Schultes et 1.
Cabrera 13898. — Rio Vaupés, between Mitu and Javarete. May 14-24, 1953.
R.E. Schultes et 1. Cabrera 1927. — Rio Vaupes, Mitu and vicinity, Urania.
‘*Flowers orange. Roots red-orange. On bare granite slope.’’ September 27 -
October 20, 1966. R.E. Schultes, R.F. Raffauf et D. Soejarto 24325.

The Kubeo medicine men on the Rio Kuduyari administer a
tea made of the crushed leaves and roots when they treat a
condition described as “‘shaking all over’’.

Nothing apparently is known of the chemistry of Schiekia.

AMARYLLIDACEAE
Eucharis amazonica Linden in Illustr. Hort. 28 (1881) 30.

178

Ecuapor: Dureno, Rio Aguarico, Napo. ‘‘Cultivated in village; also
grows in forest not far from rivers.’’ February 7, 1966. H.V. Pinkley 105.

The whole plant (including the bulb) is boiled and steeped to
prepare an emetic tea. Pinkley reports that the Kofans, who
call the plant kon-si-ahipa-cho, believe that this decoction
enables them to attain greater accuracy in using the blow gun.

Apparently no chemical investigations of Eucharis have
been published.

ARACEAE

Anthurium crassinervium (Jacg.) Schott var. caatingae R.E.

Schultes var. nov.

A Anthurio crassinervio principaliter foliis chartaceis (non
firme coriaceis), anguste lanceolatis, spathae spadicisque colo-
re differt.

CoLomstia: Comisaria del Vaupés, Rio Vaupés, Mitu and vicinity. At base
of Cerro de Mitu. ‘‘Spathe grey-purple. Spadix purple. In white sand”’.
September 27 - October 20, 1966. R. E. Schultes, R. F. Raffauf et D. Soejarto
24202 (TYPUS in Herb. Gray).

A warm decoction made of the leaves of this coriaceous
variety is employed by the Kubeo Indians of the region of Mitu
as an ear wash to relieve a condition due probably to fungal
infection.

ZINGIBERACEAE

Costus amazonicus (Loes.) Macbride in Field Mus. Publ. Bot.
11 (1931) 13.

CoLomBia: Comisaria del Vaupés, Rio Kananari, Cerro Isibukuri. Flow-
ers pale pink. Bracts green, slightly red at base on both surfaces. Height 10
feet. Taiwano name = nd-ka’. March 8, 1951. R.E. Schultes et I. Cabrera
13263.

Costus erythrocoryne K. Schumann in Engler, Pflanzenr. Zin-
gib. (1904) 410.

CoLomsia: Comisari del Vaupés. Same locality. “‘Bracts all deep scarlet
on both surfaces. Flowers red. Height 6 feet. Kabuyari name = wa'-roo-ne.
Taiwano name = fia-ka'. March 8, 1951. R. E. Schultes et I. Cabrera 13262.

Both of these species are similarly employed by the Taiwano
and Kabuyari Indians of the Rio Kananari: the leaves are dried

179

and pulverized, and the powder is snuffed into the nostrils to
staunch persistent nose-bleeding, a condition frequent among
aging individuals.

Hedychium coronarium Koernicke in Retzius, Obs. 3 (1783)

7.

COLOMBIA: Comisaria del Vaupés, Rio Kuduyari, Cerro Yapoboda.
‘Flowers white, fragrance of narcissus. Plant 6 feet tall."’ October 1, 1951.
R.E. Schultes et 1. Cabrera 14188.

The Kubeos take a decoction of the root to relieve pains in
the chest and arms. Their name of the plant is ma-tsé'-ko-ra.

ORCHIDACEAE

Dichaea muricata (Sw.) Lindley, Gen. and Sp. Orch. Pl. (1833)

209.

Ecuabor: Dureno, Rio Aguarico, Napo. October 20, 1966. H.V. Pinkley
526.

A wash is prepared from this orchid by the Kofans for
treating eye infections. The Kofan name is shahasi-sehé'-pa.

Psygmorchis pusilla (L.) Dodson et Dressler in Phytologia 24

(1972) 288.

Oncidium pusillum (L.) Reichenbach fil. in Walp. Ann. Bot.
Syst. 6 (1863) 714.

EcuaApbor: Dureno, Rio Aguarico, Napo. December 20, 1965. H.V.
Pinkley 11.

The Kofan Indians, who call this epiphyte atiipa-kashaikie-
si-sehé'-pa, treat lacerations with a wash prepared by boiling
the plant in water.

MORACEAE

Brosimum utile (HBK.) Pittier var. ovalifolium (Ducke) C.C.
Berg in Act. Bot. Neerl. 19 (1970) 328.
COLOMBIA: Comisaria del Vaupes, Rio Apaporis, Jinogoje. **Tree 40 feet
tall. | foot in diameter.”” June 15, 1952. R.E. Schultes et 1. Cabrera 16735.
The bark of this tree is extensively employed for making
clothing, especially uniforms for dancing. The latex, taken
cautiously, 1s considered a tonic.

180

The latex is employed medicinally in parts of Colombia in the
belief that it cures asthma and other pulmonary and bronchial
ailments. (Garcia-Barriga: Flora Medicinal de Colombia 1
(1974) 241).

The Makunas call the tree wa-s6-gee; the Makus, bawn'-tég;
the Yukunas, ma-ree-m ‘a.

Coussapoa cinnamomea Cuatrecasas in Caldasia 7 (1956) 288.

CoLomsia: Comisaria del Amazonas, Trapecio Amazonica, Rio Loreto-
yacu. October, 1945. R.E. Schultes 6693. — Same locality. September 1946.
R.E. Schultes et G. Black 8269.

The Tikuna Indians of the Trapecio Amazonico formerly

employed leaves and fruit of Coussapoa cinnamomea,
pounded and mixed with mud, as a fish poison.

Coussapoa magnifolia 7récul var. glabrescens Cuatrecasas var.
nov.

Differt a var. magnifolia typica, foliis supra leavibus glaber-
rimis vel spasissim pilis, subtus tandum pilis tenuibus adpressis
sparsis praeditis; stipulis subglabris; ramulis inflorescentiae
minutissime hirtulis.

BRAZIL: Estado do Amazones, Rio Negro, between Uaupes (Sao Gabriel)
and Sao Felipe. October, 1947. R. E. Schultes et J. Murca Pires 8976 (spec.
masc.) — COLOMBIA: Comisaria del Vaupés, Rio Vaupes, Raudal del

Yurupari, quartzitic base. ‘‘Tree 50 feet tall’? November 1951. R. E. Schultes
et 1. Cabrera 1958 (spec. fem.) (TYPUS in U.S.Nat. Herb.)

Coussapoa orthoneura Standley in Field Mus. Publ. Bot 17

(1947) 16).

BRAZIL: Estado do Amazonas, Rio Tikié, Taracua. January 28 - February
9, 1948. R.E. Schultes et F. Lopez 9685. — CoLcomBia: Comisaria del
Amazonas, Rio Apaporis, Soratama. ‘‘Gigantic tree 70 feet tall. Fruit yellow.
Piunave = koom-ka'.”’ August 27, 1951. R.E. Schultes et 1. Cabrera 13774. —
Comisaria del Vaupés, Rio Apaporis, Raudal de Jerijerimo, ‘Small tree.
Fruit red. On rock savannah cliff.’” February 14, 1952. R. E. Schultes et I.
Cabrera 15324. — Rio Vaupés, Raudal de Tatu. *‘ Flowers yellow. Small tree,
60 feet. Hanging over cataract’’. October 10, 1966. R.E. Schultes, R.F.
Raffauf et D. Soejarto 24379.

The Puinave Indians chew the fruits to cure mouth sores.
The leaves of Schultes, Raffauf et Soejarto 24379 gave a
positive test for alkaloids with Dragendorff reagent.

181

Coussapoa trinervia Spruce ex Mildbread in Notizbl. 10 (1928)
416.

COLOMBIA: Comisaria del Amazonas, Rio Apaporis, Soratama. *‘Tree.
Fruit green. Flood-forest’’. Aug. 16, 1951. R.E. Schultes et 1. Cabrera 13589.

The Indians of the Rio Apaporis frequently chew the hard,
astringent fruits to treat sores of the tongue and mouth.

Helicostylis scabra (Machride) C.C. Berg in Act. Bot. Neerl. 18
(1969) 464.

COLOMBIA: Comisaria del Vaupes, Rio Apaporis, Raudal Yayacopi (La
Playa). **Ernormous tree. Latex white. Fruit green, edible.”” February 18,
1952. R.E. Schultes et 1. Cabrera 15466.

The Puinave Indians call this tree han-shee-ma; the Makus,
meeé-o. The latex is applied to abrasions of the skin in the belief
that it prevents infection.

Poulsenia armata (Miq.) Standley in Trop. Woods, no. 3 (1933)

4,

CoLomsia: Comisaria del Putumayo, Rio Sucumbios, Santa Rosa. April
7-8, 1942. R.E. Schultes 3617.

The Kofan Indians employ the bark in the manufacture of
cloth for clothing and bedding.

Pourouma Schultesii Cuatrecasas in Caldasia 7 (1956) 303.
COLOMBIA: Comisaria del Vaupeés, Rio Apaporis, Jinogojé. September 25,
1952. R.E. Schultes et 1. Cabrera 17615.
The Indians of the lowermost Rio Piraparana burn the bark
of this tree and apply the ashes to running sores and ulcers.
The Puinave name is mui.

Pseudolmedia laevigata 7récu/ in Ann. Sci. Nat., ser. 3, 8 (1847)
ES.
COLOMBIA: Comisaria del Amazonas, Rio Apaporis, Soratama. ** Fruit
red, soft. Small tree.’ February 4, 1952. R.E. Schultes et 1. Cabrera 15147.
The fruits of this tree are said by the Indians of the Rio
Apaporis to be toxic.
Cyanogenesis has been reported from Pseudolmedia (Gibbs,
R.D.: Chemotaxonomy of Flowering Plants 3 (1974) 1818).

182

MyYRISTICACEAE

Virola albidiflora Ducke in Journ. Wash. Acad. Sci. 26 (1936)
259

COLOMBIA: Comisaria del Vaupés, Rio Vaupés, Mitu and vicinity. **Small
tree. Flowers yellow.’’ September 8, 1951. R.E. Schultes et 1. Cabrrea
13974.

The Kubeos call this tree po-ham-bo'-raek; the Tukanos,
sé-ne-'no. Both peoples employ the fresh bark resin in treating
skin sores.

There appears to be a rather generalized use of the bark resin
of various species of Virola to alleviate or cure fungal infec-
tions of the skin and to hasten the healing of wounds (Schultes
and Holmstedt in Lloydia 34 (1971) 61-78).

CONNARACEAE

Rourea cuspidata Bentham et Baker var. densiflora (Steyer-
mark) Forero in Mem. N. Y. Bot. Gard. 26 (1976) 48.

CoLomsia: Comisaria del Amazonas, Trapecio Amazonico, Loretoyacu
River. October 1945. R.E. Schultes 6684. Same lonality. R.E. Schultes et
G.A. Black 8305. September 1946. — Rio Amazonas, La Victoria. August 30,
1939. L. Williams 2963.

The Tikunas of the region of Leticia, Colombia, indicate that
this species is toxic, but they no longer have a use for it.

Forero (/oc. cit., 28) reports that the seeds and roots of some
species of Rourea have been used as dog poisons, source of the
Brazilian vernacular name mata-cachorro.

Rourea glabra Humboldt, Bonpland et Kunth, Nov. Gen. et
Sp. 7 (1825) 41.
A decoction of Rourea glabra is said to be employed as a
medicine for sore throat (Forero: Joc. cit., 28).

LEGUMINOSAE
Tachigalia cavipes (Spruce ex Benth.) Macbride in Field Mus.
Publ. Bot. 13, pt. 3 (1943) 127.

CoLomBIA: Comisaria del Vaupés, Mitu. December 18, 1939. E. Pérez-
Arbeldez et J. Cuatrecasas 6738. — Rio Kubiyu, Guranhuda. June 30, 1958.
H. Garcia-Barriga, R.E. Schultes et H. Blohm 16061. — Rio Kananari. Cerro

183

Isibukuri, 250 m. - 700 m. November 29-30, 1951. H. Garcia-Barriga 13779.
— Rio Piraparana. Loma Buc-chia. August 28-31, 1952. H. Garcia-Barriga
14286 — Rio Piraparana, middle course. January 9, 1952. R.E. Schultes etl.
Cabrera 17137. — Rio Vaupés, below mouth of Rio Kubiyu, March 26, 1970.
D. Soejarto et T.E. Lockwood 2430. — Right tributary of Rio Macu-Parana,
upland forest. ‘‘Ingredient of we-wit-kat-ku (‘no children medicine’) of
Bara-Maku. June 1-8, 1970. P. Silverwood - Cope 10. — Rio Vaupes, about |
km. below Mitu. **Tree 12 m. tall, spreading, inundated; leaves silvery green
witha swollenrachis at first pair of leaflets housing extremely ferocious ants;
flowers with calyx pale green, corolla yellow.”* July 3, 1976. J.L. Zarucchi
ISIOA. **Tree 15 m. tall, spreading, at edge of river, inundated. Leaves
silvery white beneath; leaves with swollen rachis at the first leaflet pair,
which house ants (these being the fiercest ants encountered by the collector
in the Vaupés). Flowers bright yellow, showy”’. June 25, 1976.J.L. Zarucchi
et M. Balick 1746.

A common plant along the banks of the rivers in the Colom-
bian Vaupes, Tachigalia cavipes has a number of medicinal
uses in the native ethnopharmacopeia. A tea of the leaves is
widely valued in the Colombian Vaupes as a febrifuge. Bark of
the stem is employed as an emetic and as a medicine when the
intestinal system has been *‘poisoned”’ by ingestion of tainted
fish. When the powdered leaves are mixed with the pulverized
leaves of a species of Combretum, they are employed as a body
insect-repellant. According to Zarucchi (Zarucchi et Balick
1746), the ‘‘bark is macerated to make a preparation used to
clean bad cuts and wounds; also, the bark is powdered and
used to ‘dry up’ cancres of the mouth and lips from over-use of
coca’.

The Kubeo Indians dry and powder the leaves to mix with
farina (flour of Manihot esculenta) when there is blood in the
stool.

The Maku Indians who live near the Barasana tribe along the
Rio Maku-Parana apparently employ Tachigalia cavipes as an
antifertility agent.

Tachigalia paniculata Auvb/et var. comosa Dwyer in Ann. Mo.
Bot. Gard. 41 (1954) 240.

CoLomBIA: Comisaria del Amazonas, Rio Loretoyacu. September 1946,
R.E. Schultes 8266. Same locality. September 1946, R.E. Schultes et H.
Garcia-Barriga 8360. — Comisaria del Vaupes, Rio Apaporis, Jinogoje.
September 3-11, 1952. H. Garcta-Barriga 14418.

The Tikuna Indians of the Leticia area of Colombia utilize

184

the seeds of Tachigalia paniculata as a medicine, preparing a
strong emetic by boiling them in water.

Tachigalia ptychophysca Spruce ex Bentham in Martius, FI.

Brasil. 15, pt. 2 (1876) 229.

CoLomBIA: Comisaria del Vaupés, Raudal de Jerijerimo and vicinity. July
7, 1951. R.E. Schultes et I. Cabrera 12955.

This species of Tachigalia is reputed to be valuable as an
aphrodisiac: the unripe pods are boiled and eaten. Schultes et
Cabrera 12955 represents apparently the second collection of
this very characteristic species of Tachigalia. It is the first from
Colombia. The type was collected by Spruce (Spruce 2022) at
Ipanore on the Rio Uaupes in Brazil.

EUPHORBIACEAE

Hevea brasiliensis (Willd ex Juss.) Mueller-Argoviensis in Lin-
naea 34 (1865) 204.

PERU: Departamento de Loreto, Provincia Maynas, Rio Ampiyacu, Pebas
and vicinity. “‘Tree 25 m. tall in secondary forest. Isolated.’ April 23, 1977.
T. Plowman, R.E. Schultes et O. Tovar (Alpha-Helix Amazon Expedition,
1976-1977, Phase VII) 6995.

Hevea guianensis Aublet, Hist. Plant. Guian. Frang. 2 (1775)
871.

PERU: Departamento de Loreto, Provincia Maynas, Rio Ampiyacu. Pucu
Urquillo and vicinity (near Pebas). ‘*Tree 60 ft. tall, 10 inches diameter. Bark
smooth, tan-grey. Latex cream.”’ April 28, 1977. 7. Plowman, R.E. Schultes
et O. Tovar (Alpha-Helix Amazon Expedition, 1976-1977, Phase VII) 7/25.

In view of the report in the literature (Raffauf, R. F, A
Handbook of Alkaloids and Alkaloid-containing Plants (1970))
of several alkaloids from the rubber tree, Hevea brasiliensis
(presumably from cultivated material collected in Malaysia),
Phase VII of the Alpha-Helix Amazon Expedition 1976-1977
carried out analyses of the two species, H. brasiliensis and H.
guianensis, growing in the region of Pebas, Peru, under natural
conditions. Both species were found to be alkaloid-negative.

Micrandra Spruceana (Baill.) R.E. Schultes in Bot. Mus.
Leafl., Harvard Univ. 15 (1952) 217.

185

PERU: Departamento de Loreto, Provincia Maynas, Rio Ampiyacu, Puca
Urquillo and vicinity. *‘Tree 65 m. tall with large buttresses, forming inter-
woven props at base. Growing in upland primary forest. Latex white, sparse.
Fruit green. Seeds brilliant, smooth, red-brown.”’ April 22, 1977. T. Plow-
man, R.E. Schultes et O. Tovar 6951. — Pebas and vicinity. Trail north from
town. ‘Fruits and seeds found on ground under large buttressed tree in
chacara.”’ April 10, 1977. T. Plowman, R. E. Schultes et O. Tovar (Alpha-
Helix Amazon Expedition 1976-1977, Phase VII) 6735.

Notwithstanding intensive study of Micrandra (including
Cunuria) over a large area of the northwest Amazon, I have not
encountered a medicinal use of the plant, until recent research
amongst the Witotos and Boras now residing in the vicinity of
Pebas in Amazonian Peru. Both Indian tribes employ the
sparse latex of this tree, which they recognize as “‘cousin”’ of
Hevea, known by the Peruvian name of conoco, for treating
sores of the gums and mucous membranes of the mouth and to
staunch the flow of blood following severance of the umbilical
cord. Upon questioning, the Indians assured me that not any
latex—i.e., latex from Hevea (which is locally much more
abundant) and from the numerous moraceous trees of the area
— would serve the same purpose, since they were more caustic
and failed to effect rapid congealment of the blood.

STERCULIACEAE

Theobroma grandiflorum ( Wil/d.) K. Schumann in Martius, FI.
Bras. 12, pt. 3 (1886) 76, t. 17.

COLOMBIA: Comisaria del Amazonas, Rio Amazonas, near mouth of Rio
Loretoyacu and Puerto Narino. *‘Tree. Cultivated. Leaves pale beneath.
Staminodes deep red. Alkaloid-negative.”’ September 13-15, 1966. R.E.
Schultes, R.P. Raffauf et D. Soejarto 24165.

PERU: Departamento de Loreto, Rio Amazonas, Iquitos. Fruit purchased
in market. May 7, 1977. T. Plowman, R.E. Schultes et O. Tovar 7301.

During Phase VII of the Alpha-Helix Amazon Expedition
1976-1977, an examination of the seeds of Theobroma grandi-
florum — the cupuassu of Brazil — was carried out. Gas
chromatography - mass spectrometry indicated a single com-
pound which was identified as tetramethyluric acid, probably a
new alkaloid.

In view of this analysis, it is of interest that a spot test with
Dragendorff reagent on Theobroma grandiflorum leaves

186

(Schultes, Raffauf et Soejarto 24165) gave a negative result for
alkaloids.

Theobroma subincanum Martius in Buchner, Repert. Pharm.
35 (1830) 23.

BRAZIL: Estado do Amazonas, Rio Purus, Rio Apitua, vicinity of
Jamamadi Indian village. Forest on terra firma. “*Tree, 15 m. x 15 cm.
diameter. Bark used in snuff. The bark is stripped, burnt and the ash mixed
with tobacco (13929) to produce a narcotic snuff. Cowadimani.”” July 1, 1971.
G.T. Prance et al. 13933. — Same locality and date. “Seedlings of tree
cultivated by Jamamadi Indians as an alternative to Theobroma (13933) in
making their narcotic snuff. Bark is burned and the ashes are added to
tobacco (13928)"’. Cupiit. G.T. Prance et al. 13939. — Rio Cunhua at Deni
Indian village. ‘Forest beside stream. Tree, 12 m. x 15 cm. diameter. The
bark ash used as an ingredient of Deni Indian snuff; the fruit eaten by Deni
Indians. Mapanaha (Deni), cupui (Port.)’’. November 29, 1970. G.T. Prance
etal. 16515. — Rio Purus, Jamamadi Mission Station, 30 km. west of Labrea.
Forest on terra firma. ‘‘Treelet, 3 m. tall. Bark ash mixed with tobacco leaves
as components of Jamamadi snuff. Shina (Jamamadi)’’. June 28, 1974. D.G.
Campbell et al. P21258.

CoLomBiA: Comisaria del Vaupés, Rio Piraparana. *‘Cultivated tree. Bark
pounded and burned to mix with tobacco and yakee snuff.’ August 24, 1952.
R.E. Schultes et 1. Cabrera 17005.

There may well be a reason for the choice of bark of this
species of Theobroma for the alkaline ash mixture in tobacco
and Virola snuffs over such a wide area of the Amazon. The
Indians of the Rio Piraparana region of the Colombian Vaupés
prefer this ash to mix with the dried and powdered Virola resin.
Natives in other parts of the Colombian Amazon (e.g., the
Yukunas of the Rio Miritiparana) use it to make ashes for
tobacco-snuff. The several reports by Prance and his collabor-
ators are based on specimens from the Rio Purus of Brazil, an
area some 600 air-miles from the Colombian locality cited
above and from totally unrelated Indians. There are likewise
reports, but without voucher specimens, of the employment of
the bark-ashes of Theobroma for this purpose from numerous
other Amazonian localities.

THY MELAEACEAE

Schoenobiblus peruvianus Stand/ey in Field Mus. Publ. Bot. 6
(1936) 169.

CoLomBIA: Comisaria del Amazonas, Rio Loretoyacu. October, 1946.

187

R.E. Schultes et H. Garcia-Barriga 8418, — Comisaria del Putumayo, Rio
Guamues, Santa Rosa. September 6, 1966. H.V. Pinkley 436. — Rio San
Miguel (Sucumbios), Santa Rosa. November 30, 1966. H.V. Pinkley 565.

Among the Tikuna Indians of the Leticia area of Colombia,
the dried leaves of this shrub are powdered and made into a
poultice to help the healing process in cases of persistent and
infected cuts or wounds. The Tikunas know the plant as ka-
we -re-ta. Schoenobiblus peruvianus is utilized by the Kofan
Indians in preparing a kind of curare (Schultes in Bot. Mus.
Leafl., Harvard Univ. 13 (1949) 285; Pinkley 565) and as a fish
poison (Pinkley 436). Chemical studies of Schoenobiblus ap-
parently have not been published.

UMBELLIFERAE

Niphogeton ternatus (Willd. ex R. et S.) Mathias et Constance
in Univ. Cal. Publ. Bot. 23 (1951) 409.

CoLoMBIA: Comisaria del Putumayo, road from Sibundoy to Pasto, bet-
ween La Marta and Paramo de San Antonio. Alt. 2900-3180 m. ‘Tea of leaves
used for colic in Sibundoy Valley.” June 1, 1946. R.E. Schultes et M.
Villarreal 7814,

This plant is highly esteemed by the Kamsa Indians of Si-
bundoy for the treatment of digestive disorders. A decoction of
the leaves is taken. There appears to have been no phytochem-
ical study pointing to active principles in this species.

STYRACACEAE

Styrax Tessmannii Perkins in Notizbl. 10 (1928) 459.
COLOMBIA: Comisaria del Amazonas, Rio Apaporis, Soratama. March 26,
1952. R.E. Schultes et 1. Cabrera 16047. — Trapecio Amazonico, Rio
Amazonas, Rio Loretoyacu. R.E. Schultes 7/144.
The leaves of Styrax Tessmannii are commonly applied
crushed to fungal infections between the toes. This use is
common amongst several tribes of the middle Rio Apaporis.

Styrax yapobodensis (J. /drobo et R.E. Schultes) Steyermark in
Fieldiana 28 (1953) 492.

COLOMBIA: Comisaria del Vaupes, Rio Kuduyari, Yapoboda. *‘Golden
pubescence on under side of leaf. Height 15 feet. Flowers white.’ October
5-6, 1951. R.E. Schultes etl. Cabrera 14394. — Same locality. **Small treelet

188

up to 8 feet.”’ April 1953. R.E. Schultes et 1. Cabrera 20012. — Rio Kubiyu,
Cerro Kanenda. Savannah. ‘‘Bush. Height 9 feet. Under part of leaf sordid.
Flowers white; anthers yellow’. November 10, 1952. R.E. Schultes et I.
Cabrera 18311.

The Kubeo Indians crush the leaves of this small shrub to
apply to warts.

POTALIACEAE

Potalia Amara Aublet, Hist. Plant. Guian. Frang. 1 (1775) 394,
on

BRAZIL: Estado do Amazonas, Rio Uneiuxi, Maku Indian village, 300 km.
above mouth. ‘Forest on terra firma. Treelet, 3 m. tall. Root bark scraped,
mixed with Menispermaceae /5560 as ingredient for Maku snake-bite cure.
Awuibiden: aw = snake (Maku)’’. October 23, 1971. G.T. Prance, P.J.M.
Maas, D.B. Woolcott, O.P. Monteiro et J.F. Ramos 15559.

CoLomsia: Comisaria del Vaupés, Rio Vaupés, Mitu and vicinity. *‘Mar-
tiguaje’’. September 16, 1939. J. Cuatrecasas 6852. — Same locality. Sep-
tember 8, 1951. R.E. Schultes et I. Cabrera 13972. — Comisaria del
Amazonas, Rio Apaporis, Soratama. ‘‘Shrub 2 m. tall. Flowers yellowish
green. Leaves paler green beneath, dark green above. Puinave name =
chin-wee'. Growing in flood forest’’. August 27, 1951. R.E. Schultes et 1.
Cabrera 13726. — Comisaria del Vaupés, Rio Vaupés, Cerro de Mitu.
‘‘Flowers yellow. Used against snake-bite. Height 5 ft. Martiguaje’’. August
20, 1960. R.E. Schultes 22718.

PERU: Departamento de Loreto, Provincia Maynas. Mishuyacu, road to
Quistococha, near Iquitos. ‘‘Curarina’’. October 15, 1965. J. Torres 109, —
Rio Napo, Negro Urco. ‘‘Curarina (sp.); cu’ yacono’ -le. Treelet, 2 m.”°.
August 27, 1966. R.T. Martin et C.A. Lau-Cam 1312. — Millpa Canio.
“Curarina-sacha’’. March 26, 1968. F. Tina et M. Tello (Amazon Natural
Drug Co.) 2074. — Provincia Mariscal Caceres, Tocache Nuevo.
““Curarina-sacha. Los nativos tomen las hojas en infusion para picadura de
viboras’’. November 10, 1969. J. Schunke V. 3610. — Rio Yaguasyacu,
affluent of Rio Ampuyacu. Brillo Nuevo and vicinity. ** Bora = okaji-kahpuu.
Fresh leaves chopped with water. Taken internally for snake-bite or against
any poisonous animal (raya, isula). Calms the body and eliminates pain’’.
April 12, 1977. T. Plowman, R.E. Schultes et O. Tovar (Alpha-Helix Amazon
Expedition 1976-1977, Phase VII) 6803. — Same locality. ‘**Unbranched
treelet, 4 m. tall, in disturbed upland forest. Leaves coriaceous, shiny.
Rhachis and inflorescence yellow. Fruit green. Alkaloid-negative’’. April 15,
1977. T. Plowman, R.E. Schultes et O. Tovar 6895.

It is obvious that in the northwest Amazon Potalia Amara is
considered a valuable and very versatile remedy. The Maku
Indians of Brazil mix the bark scrapings with the bark of a
menispermaceous plant to treat snake-bite. The several tribes
of the Colombian Vaupés likewise consider it efficaceous for

189

snake-bite. This same use prevails also in Amazonian Peru and
Brazil.

Although the plant has a wide reputation as a treatment for
snake-bite, the Indians of the Vaupés in Colombia appear to
value it primarily in infusion as an emetic to be used in cases of
food poisoning, frequent during tribal festivals. The Bora In-
dians in Peru take the leaves finely chopped in water ‘‘to calm
the body and eliminate pain’’, not only in the case of snake-
bite, but for the stings of the tail of the fresh water skate (raya)
and of large, poisonous ants (isula).

It is difficult to understand the paucity of chemical investiga-
tion of a plant ethnopharmacologically so important over so
wide an area. Fresh material (Plowman, Schultes et Tovar
6803) was chemically examined on the Alpha-Helix Amazon
Expedition 1976-1977: *‘Only squalene and methyl esters of
fatty acids were found . . . No alkaloids were present’’. Squa-
lene is widely distributed in plants, known from 14 families of
higher plants and from yeasts.

Known as femblador in Venezuela, an infusion of the bark is
employed as a laxative (L. Williams 14352). In the western
Amazon of Brazil, where the plant is called anabi and pao de
cobra (‘snake plant’’), Potalia Amara is recognized as toxic.
The leaves and shoots are made into a bitter infusion employed
in treating syphilis, and a decoction of the leaves is used as a
wash for eye infections. A tea of the leaves is emetic in large
doses and is taken to empty the stomach of tainted cassava
flour. Folk medicine in Brazil maintains that the root is valu-
able in treating snake-bite (Le Cointe, P.: Amazonia Brasileira
3 (1943) 20.

In Peru, Potalia Amara is known as sacha-mangua (L.
Williams in Field Mus. Nat. Hist. Bot. Ser. 15 (1936) 418),
curarina and curarina-sacha (‘curanina’’ is a name widely
employed in Hispanic America for snake-bite potions). The
most widely known term for the plant in Colombia is mar-
tiguaje. One of the Brazilian names, pao de cobra, referes to
the widespread esteem that the plant enjoys as an antidote for
snake venom.

It would appear that a plant of such varied therapeutic uses
should be the subject of intense phytochemical study.

190

SOLANACEAE

Cyphomandra crassifolia (Ort. ) Macbride in Publ. Field Mus.
Nat. Hist., Bot. 8 (1930) 112.

CoLomBiA: Comisaria del Putumayo, Rio Sucumbios, Conejo and vicin-
ity. Altitude 300 m. ‘Bush. Flowers green, anthers white. Fruits round,
green, hard.’ April 2-5, 1942. R.E. Schultes 3652. — Puerto Ospina and
vicinity. March 25-26, 1953. R.E. Schultes et 1. Cabrera 18979. — Comisaria
del Amazonas, Rio Amazonas, Leticia. September 20, 1945. R.E. Schultes
6541. — Comisaria del Vaupés, Rio Apaporis, Soratama. August 17, 1951.
R.E. Schultes et I. Cabrera 13044. — Rio Vaupés, Mitu and vicinity. Sep-
tember 7, 1951. R.E. Schultes et I. Cabrera 13903.

The Kofan Indians employ the leaves of Cyphomandra cras-
sifolia for dying clay pots black. A tea of the leaves is used by
the Indians in the Mitt region to expel intestinal parasites.

Scopoletin has been isolated from Cyphomandra
(Hegenauer, R.: Chemotaxonomie der Pflanzen 6 (1973) 418.
Tannins and tetraterpenes have likewise been reported from
the genus (Gibbs loc. cit., 3 (19746 1762-1765).

Cyphomandra dolichorachis Bitter in Fedde Rep. 17 (1921)

350.

CoLomBIA: Comisaria del Putumayo, Sibundoy. Altitude 2225-2300 m.
May 29, 1946. R.E. Schultes et M. Villarreal 7650.

The Kamsa Indian medicine men employ a decoction of the
leaves as a vermifuge.

Cyphomandra endopogon Bitter in Engler, Jahrb. 54, Beibl.
119 (1916) 16.

CoLomBIA: Comisaria del Amazonas, Rio Loretoyacu. August 19, 1964.
A. Ferndndez-Pérez 6863. — Same locality. ‘‘Small tree to 6 m. tall in
secondary growth. Heliophile. Corolla green; stamens white at anthesis.
Fruit a berry, ovoid, yellow at maturity. Whole plant gives strong narcotic
odor’’. January 28 - February 7, 1969. 7. Plowman, T. Lockwood, H. Ken-
nedy et R.E. Schultes 2332.

A spot test with Dragendorff reagent indicated that this
species (Ferndndez 6863) is weakly alkaloidal.

Juanulloa ochracea Cuatrecasas in Brittonia 10 (1958) 148.

CoLomsia: Comisaria del Vaupés, Rio Apaporis, Raudal Yayacopi (La
Playa) ‘‘Bush. Flowers green. Epiphyte (?)."° February 16, 1952. R.E.

19]

Schultes et 1. Cabrera 15412. — Comisaria del Putumayo, Rio Putumayo,
Puerto Ospina. **Bush 15 feet tall. Flowers yellowish; bracts orange. Leaves
crassulent.’’ March 23-26, 1953. R.E. Schultes et 1. Cabrera 18960. — Com-
isaria del Caqueta, Rio Caqueta, Floresta, c. 15 km. down from Puerto
Limon. **Climbing epiphyte shrub on fallen tree, 3 m. tall. Calyx red: corolla
yellow. Leaves coriaceous.’’ December 20, 1968. 7. Plowman 2176.

The inhabitants in the vicinity of Puerto Limon, who call this
plant ayahuasca, employ the leaves and the trunk to treat
wounds. The vernacular name — commonly applied to an
hallucinogenic plant (Banisteriopsis Caapi) and a preparation
from it — suggests its possible use as an intoxicant (Schultes in
Bot. Mus. Leafl., Harvard Univ. 23 (1972) 140).

The alkaloid parquine has been reported from this genus
(Raffauf, loc. cit.)

Markea coccinea L.C. Richard in Act. Soc. Hist. Nat. Paris 1
(1792) 107.

COLOMBIA: Comisaria del Vaupes, Rio Apaporis, Raudal de Jerijerimo.
July 8, 1951. R.E. Schultes etl. Cabrera 13441. — Rio Pacoa. ** Vine. Flowers
red.’’ February 7-12, 1952. R.E. Schultes et I. Cabrera 15251. — Rio
Apaporis, Soratama. **Medium sized treelet. Flowers white.’’ January 1952.
R.E. Schultes et 1. Cabrera 19846. — Rio Vaupes, between Mitt and
Javarete, Cerro de Tipiaca. *‘Climber. Flowers vermillion-orange.’> May
14-15, 1953. R.E. Schultes et 1. Cabrera 19305.

The Desano Indians, who call the plant see-nan-da °, apply a
decoction of the leaves of this relatively rare species to treat

conjuctivitis and other eye affections.
Nothing apparently is known of the chemistry of Markea.

Solanum albidum Duna/, Hist. Solan. (1813) 206.

PERU: Departamento de Loreto, Pucallpa. Altitude 200 m. ‘'In low forest.
Flowers lilac. Plant 2-3 m. Vulgar name: mullaca. Las hojas las utilizan los
indios Shipibos para labado vaginal en enfusion. El tronco tiene espinas.”’
November 20, 1965. J. Schunke V. 1014.

An infusion of the leaves is reported as a vaginal wash
amongst the Shipibo Indians of Peru.

Solanum apaporanum R.E. Schultes in Bot. Mus. Leafl., Har-
vard Univ. 13 (1949) 292.

COLOMBIA: Comisaria del Vaupes, Rio Vaupes, Mitu and vicinity. ** Vine.
Spines on stem. Flowers white. Fruit orange. Alkaloid-positive.’’ September
27 - October 20, 1966. R.E. Schultes, R.F. Raffauf et D. Soejarto 24300.

7?

The abundant fruits of Solanum apaporanum are dried and
powdered by the Kubeo Indians who apply the dust to the body
at night as an insect repellant.

A Dragendorff reagent spot test on fresh material indicated
that this species is alkaloid positive.

Solanum campaniforme Roemer et Schultes, Syst. 4 (1819)

662.

CoLoMBIA: Comisaria del Vaupés, Rio Vaupés, Mitu and vicinity. “*Small
tree, 10 feet tall. Petals deep blue; fruits green. In secondary growth. Alka-
loid-positive.’’ September 27 - October 20, 1966. R.E. Schultes, R.F. Raffauf
et D. Soejarto 24292.

Desano Indians boil the leaves to make a strong infusion in
which they bathe feet badly infected with niguas or sand-fleas
before extracting the egg-sacs with spines.

A spot test with Dragendorff reagent indicated the presence
of alkaloids in the leaves.

Solanum jamaicense Miller, Gard. Dict., ed. 8 (1768) no. 17.
CoLomBiA: Comisaria del Amazonas, Leticia and vicinity. “‘Flowers
white. Shrub.’ August - September, 1966. R.E. Schultes, R.F. Raffauf et D.
Soejarto 24098.
An alkaloid test with Dragendorff reagent indicated that this
species is doubtfully alkaloidal.

Solanum mammosum Linnaeus, Sp. Pl. (1753) 187.

CoLomsia: Comisaria del Putumayo, Rio Putumayo, between Puerto Asis
and Puerto Ospina. ‘‘Dooryard; not planted. Shrub 1.75 m. Corolla purple.
Fruit bright yellow, pulp white, seeds reddish brown. Fruit used as a cock-
roach poison. Cocona, cucunu, resgalgal, chufchu.”’ May 16, 1963. M.L.
Bristol 1011. — Region of Mocoa, camino viejo Pepino - Mocoa. Altitude
500-800 m. ‘‘Regargar. Sirve para matar las cucarachas.’’ August 28, 1963.
P. Juajibioy Chindoy 266.

Ecuapor: Napo, Dureno, Rio Aguarico. February 28, 1966. “Kofan =

koko'-no-cho.”’ H.V. Pinkley 172.

Peru: Departamento de Loreto, Rio Napo, Negro Urca. ‘Cultivated shrub.

Cocona venenosa.”’ August 16, 1966. R.T. Martin et C.A. Lau - Cam 1288.
The efficacy of the powdered fruit as a specific poison for

cockroaches is widely recognized, but use of the plant for this

purpose seems nowhere to be more extensive than in the

193

westernmost parts of the Amazon amongst the Indians of Col-
ombia, Ecuador and Peru.

The collection Pinkley 1/72 reports an interesting use of this
toxic plant: “‘As a pacifier for small children.”

Solanum subinerme Jacquin, Enum. Pl. Carib. (1760) 15.
COLOMBIA: Comisaria del Vaupes, Rio Vaupés, Mitu and vicinity.
‘Height 4-5 feet. Flowers blue-purple. Weed.’ September 27 - October 20,
1966. R.E. Schultes, R.F. Raffauf et D. Soejarto 24244.
A spot test in the field with Dragendorff reagent indicated
that this species is alklaoid-positive.

Solanum Topiro Humboldt et Bonpland in Dunal, Sol. Gen.

Aff. Sym. (1816) 10.

COLOMBIA: Comisaria del Vaupes, Rio Apaporis, between Rios Kana-
nari and Pacoa. *‘Flowers greenish, anthers yellow. Bush. Fruit green,
turning orange. Kubeo = be-ta'-ka; Taiwano = dé-twa'; Tatuya = da-twa'.
Cultiivated.”” September 1, 1951. R.E. Schultes et I. Cabrera 13842.

The small seeds of this edible fruit are often dried and pul-
verized for medicinal use: amongst the Taiwano, the powder
is kept and added to powder to Erythroxylon Coca when the
mucous membranes and tongue become irritated from over-
use of coca.

Solanum verbascifolium Linnaeus, Sp. Pl. (1753) 184.

CoLomBIA: Comisaria del Amazonas, Rio Amazonas, near mouth of
Rio Loretoyacu and Puerto Narino. ‘‘Bush, 8 feet. Flowers white or pale
violet. Alkaloid-positive.’” September 13-15, 1966. R.E. Schultes, R.F.
Raffauf et D. Soejarto 24123. — Comisaria del Vaupés, Mitu and vicinity.
“Herb up to 3 feet. Flowers white, leaves used for washing. Leaves as-
perous, causing rash and itching. In secondary growth.’’ September 27 -
October 20, 1966. R.E. Schultes, R.F. Raffauf et D. Soejarto 24228.

This bushy species has long been employed in the Vaupés
as one of the most easily available plants for washing cloth-
es. It is not cultivated but is found growing spontaneously in
every house site or Manihot field. The asperous leaves cause
rashes and itching if handled over a long period.

The leaves give a positive test for alkaloid with Dragen-
dorff reagent. They undoubtedly have a high concentration
also of saponins.

194

RUBIACEAE

Isertia rosea Spruce in Martius, Fl. Bras. 6, pt. 6 (1889) 284.

CoLomBIiA: Comisaria del Vaupés, Rio Kananari, Cachivera de Palito.
July 25, 1951. R.E. Schultes et 1. Cabrera 13148.

The Taiwano Indians of the Rio Kananari value a decoction
prepared from the leaves of this plant as an effective febrifuge.
It is taken hot in quantity every several hours to reduce fever
by stimulating perspiration.

Pagamea coriacea Spruce ex Bentham in Journ. Linn. Soc. |

(1857) 110.

CoLoMBIA: Comisaria del Vaupés, Rio Apaporis, Raudal de Jerijerimo.
March, 1951. R.E. Schultes 12108. — Same locality. June 13, 1951, R.E.
Schultes et 1. Cabrera 12467; January 21, 1952, R.E. Schultes et 1. Cabrera
14953; March 15, 1952, R.E. Schultes et I. Cabrera 1593]. — Rio Karuru,
Mesa de Yambi, Savannah Goo-ran-hoo-da’, April 15-16, 1953, R.E.
Schultes et 1. Cabrera 19169.

The Indians in the region of Raudal de Jerijerimo on the Rio
Apaporis value this plant as a very efficaceous remedy for
reestablishment of the ability to walk following attacks which,
in age, appear to deprive Indians from free use of the legs. The
causes of this curious disease (which is not uncommon) are
unknown.

The bark of the young branches is scraped and, in the fresh
condition, is boiled into a decoction, which must be drunk over
a period of two or three weeks. Administration of this tea is
reputed to result in strong stimulation of the afflicted patient
and sometimes in ability to regain use of the legs.

Pagamea macrophylla Spruce ex Bentham in Journ. Linn.

Soc. 1 (1857) 110.

CoLomBIA: Comisaria del Vaupés, Rio Piraparana, Cano Paca. Septem-
ber 18, 1952. R.E. Schultes et I. Cabrera 17581.

The leaves of Pagamea macrophylla are sought by medicine
men of the Makuna tribe. They are dried, pulverized and aspi-
rated as a powder during diviniation ceremonies. There is no
indication, however, that they have hallucinogenic properties.
The Maku name of this plant in the Rio Piraparana is ma-ha-
shu'-ke-ma.

195

Retiniphyllum concolor (Spruce ex Benth.) Mueller-Argovien-
sis in Martius, Fl. Bras. 6, pt. 5 (1881) 8.

CoLoMBIA: Comisaria del Vaupes, Rio Apaporis, Raudal de Jerijerimo.
March 1951. R.E. Schultes 12110. — Rio Guainia, Puerto Colombia. October
31 - November 2, 1952. R.E. Schultes, R.E.D. Baker et 1. Cabrera 18156 —
Rio Negro, San Felipe. *‘Bush. Fruit red. Flowers green-white. °° November
2, 1952. R.E. Schultes, R.E.D. Baker et 1. Cabrera 18244. — Rio Guainia,
Cano del Caribe. **Fruit red.” November 2, 1952, R.E. Schultes, R.E.D.
Baker et I. Cabrera 1825S.

This species of Retiniphyllum in the northwest Amazon is
believed to have antituberculous properties. The leaves are
burned, and the aromatic smoke 1s allowed to permeat the part
of the round house where the patient resides. It is thought to

have purifying and therapeutic effects.

Retiniphyllum pilosum (Spruce ex Benth.) Mueller-Argoviensis
in Martius FI. Bras. 6, pt. 5 (1881) 7.

BRAZIL: Estado do Amazonas, Rio Negro, Jucabi (at mouth of Rio Curicur-
iari). ‘Bush. Flowers white. Fruit red.”’ January 17, 1948. R.E. Schultes et F.
Lopez 9633.

COLOMBIA: Comisaria del Vaupés, Rio Guainia, Puerto Colombia (opposite
Maroa). **Bush 4-5 feet. Fruit red.’ October 31 - November 2, 1952. R.E.
Schultes, R.E.D. Baker etl. Cabrera 18155. — Rio Guainia, Raudal del Sapo,
below Puerto Colombia. October 31 - November 2, 1952. R.E. Schultes,
R.E.D. Baker et 1. Cabrera 18202. — Same locality and date. R.E. Schultes,
R.E.D. Baker et 1. Cabrera 18237.

The natives who live along the Rio Guainia dry the fruit of
Retiniphyllum pilosum for medicinal use during the year as an

anthelmintic.

Retiniphyllum Schomburgkii Mueller-Argoviensis in Martius,
Fl. Bras. 6, pt. 5 (1881) 12.
COLOMBIA: Comisaria del Vaupes, Rio Negro, San Felipe. October 31 -
November 2, 1952. R.E. Schultes, R.E.D. Baker et I. Cabrera 17997.
The fruits of Retiniphyllum Schomburgkii are considered to
be effective as a vermifuge amongst the Indians of the upper
Rio Negro area and the Rio Guainia.

Retiniphyllum speciosum (Bentham) Mueller-Argoviensis in
Martius, Fl. Bras. 6, pt. 5 (1881) 10.

196

BRAZIL: Estado do Amazonas, Rio Uaupés, Serra Wabeesee, below Bela
Vista. November 17, 1947. R.E. Schultes et J. Murca Pires 9139.

CoLomBIA: Comisaria del Vaupés, Rio Negro, San Felipe. ‘‘Small tree or
large bush. Fruit red. Flowers white and red.” December 12, 1947. RAE.
Schultes et F. Lopez 9327.

The fruit of Retiniphyllum speciosum is considered by the
Kuripakos of the Rio Negro to be an efficient vermifuge when
eaten in quantity.

Retiniphyllum truncatum Mueller-Argoviensis in Martius, FI.
Bras. 6, pt. 5 (1881) I1.

CoLomBIA: Comisaria del Vaupés, Rio Apaporis, Raudal de Jerijerimo.
‘Fruit red, edible. Small bush.’’ January 13, 1951. R.E. Schultes et I. Cabr-
era 12464. — Same locality. ‘‘Bush up to 5 feet tall. Leaves and petioles
sticky when slightly dried. Flowers white. Floral axis bright red.”’ March
1951.R.E. Schultes 12111. —Same locality. June 13, 1951. R.E. Schultes etl.
Cabrera 12453. — Same locality. *‘Bush, 4 feet tall. Flowers white, red at
centre. Fruit scarlet."” September 16, 1951. R.E. Schultes et 1. Cabrera
14009. — Same locality. November 27, 1951. R.E. Schultes et 1. Cabrera
14628. — Rio Guainia, Puerto Colombia. October 31 - November 2, 1952.
R.E. Schultes, R.E.D. Baker et I. Cabrera 18170 — Rio Kubiyu, Cerra
Kanends. November 10, 1952. R.E. Schultes et 1. Cabrera 18366. — Rio
Parana Pichuna. June 1953. Schultes et Cabrera 19943. — Comisaria del
Amazonas, Rio Caqueta, La Pedrera, Cerro de La Pedrera. October 2, 1952.
R.E. Schultes et 1. Cabrera 17681.

The Taiwanos, who live near the Raudal de Jerijerimo, value
the resin of this bush for treating hemorrhoids and other causes
of rectal bleeding, applying the resin to the affected area with
the finger over a period of several days. The resin must be
taken from the fresh plant. The Taiwano name for Retiniphyl-
lum truncatum is bov-fee’.

197

BOTANICAL MUSEUM LEAFLETS VoL. 26, No. 5
May 30, 1978

COCAINE IN BLOOD OF COCA CHEWERS

seo

AND T. PLOWMAN***

Although the non-medical use of cocaine, either by sniffing
or injection, is considered harmful by many medical authori-
ties, there still is controversy as to whether the chewing of coca
leaves, as practised by South American Indians, is detrimental
or not. Experimental evidence on coca chewing gathered sci-
entifically in the field has not previously been substantiated by
measurements of blood levels of cocaine.

During Phase VII of the Alpha-Helix Amazon Expedition
1976-1977, we had occasion to study two methods of adminis-
tration of coca leaves and to determine the amount of cocaine
in blood versus time by an unequivocal method of analysis.
Whole coca leaves (Erythroxylum Coca Lam.) were obtained
from Pisac, Department of Cuzco, Peru. Coca powder (pul-
verized leaves of E. Coca mixed with Cecropia leaf ash) was
prepared by Witoto Indians of the Rio Ampiyacu, Department
of Loreto, Peru, according to the custom of the region.

Coca leaves and powder (5-10 g.) were taken orally by
human subjects in the same way that South American natives
do. The cocaine, as measured by mass fragmentography, per-
sisted in the plasma for more than seven hours and reached
concentrations from 10 to 150 ng/ml at 0.38 to 1.95 hours.
Half-lives of the elimination of cocaine have been calculated
ranging from 1.0 to 1.9 hours. The absorption half-lives ranged
from 0.2 to 0.6 hours (see Table 1).

The stimulating effect obtained seems to be well correlated

*Department of Toxicology, Swedish Medical Research Council, Karolinska Institu-
tet, S-104 01, Stockholm, Sweden.

**Institute of Plant Biology and Physiology, University of Lausanne, CH-1005 Laus-
anne, Switzerland.

*** Atkins Research Fellow in Economic Botany, Botanical Museum of Harvard
University.

199

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200

with the rising concentrations of cocaine in the blood. The
shape of the curves fits with the subjective effects reported.
The differences in stimulation between using whole coca
leaves or coca powder and taking cocaine by local application
in the nose or by intravenous injections seems to be essentially
a difference in means of administration and dosage. There is,
consequently, no reason to believe that the stimulating effect
achieved by the use of either coca leaves or powder is not due
to cocaine.

An extended report on this experimental project, of which
the present article is an abstract, will be published in the
Journal of Ethnopharmacology.

The research upon which this abstract is based was carried
out under NSF Grant No. DEV 72-02536-Garey 6-446406-
21412.

201

BOTANICAL MUSEUM LEAFLETS VoL. 26, No. 5
May 30, 1978

CHROMOSOME NUMBERS IN NEOTROPICAL
ERYTHROXYLUM (ERYTHROXYLACEAE)

T. PLOWMAN !, L. RUDENBERG2, AND C.W. GREENE?

The genus Erythroxylum includes some 200 species, the
great majority of which are found in the American tropics.
Within this genus are found several economically important
plants, most notably E. Coca Lam. and FE. novogranatense
(Morris) Hieron. These two closely related species native to
South America are extensively cultivated as the sole source of
the alkaloid cocaine. Numerous cultivars and local races of
these species occur in the Amazon and Andes, the result of at
least 6000 years of coca cultivation. The variation and evolu-
tion of coca under domestication has been little studied in
recent years and little is known about the cytogenetics and
breeding relationships of this important crop.

As part of a multidisciplinary project on coca, we have
conducted a preliminary cytological study on Erythroxylum
species. Until now, there have been no published chromosome
counts with voucher specimens of any neotropical Erythroxy-
lum; nor are any vouchered counts known for the cultivated
species. Species of Erythroxylum are notoriously difficult to
distinguish morphologically, and much confusion exists in the
taxonomic placement of many species. It is therefore essential
that herbarium vouchers accompany all chromosome studies
and that chemical analyses carried out on the genus to insure
present and future identifications.

Five chromosome counts of Erythroxylum species have been
published in the past. These are summarized in Table One.
Only one of these counts (E. Kunthianum Wall.) bears a
voucher specimen. The correct identity of the two plants de-
termined as E. Coca is especially open to question. Several

'Botanical Museum of Harvard University.
2Gray Herbarium, Harvard University.

203

related species, both wild and cultivated, are frequently mis-
identified as FE. Coca.

MATERIALS AND METHODS

Mitotic counts were made from root tip meristems from
greenhouse reared plants, pretreated in 0.004 M 8-hydroxy-
quinoline at 15°C. for 3 hours (Tijo & Levan, 1950). Root tips
were then rinsed in distilled water, stained 1/2 hour in 1% acetic
orcein: | N HC1 (9:1) with gentle heating, and squashed in 45%
acetic acid.

Meiotic studies were conducted with flower buds fixed and
stored in Carnoy’s solution (ethanol: acetic acid, 3:1). Tissue
stored in Carnoy’s for up to 4 years provided adequately pre-
served material for study. Microsporocytes were squashed and
stained in acetocarmine. Photographs were taken with a Zeiss
oil immersion lens under phase illumination. Herbarium
vouchers are preserved at the Economic Herbarium of Oakes
Ames (ECON).

RESULTS AND DISCUSSION

Chromosome numbers are summarized in Table Two. All
counts showed 2n = 24 or n = 12, in agreement with earlier
published counts for Erythroxylum. It would seem that n = 12
is the base number for the genus. Some chromosomal ir-
regularities, including chromosome bridges, were observed in
microsporocytes of E. Coca (Plowman 6165, 6183). A more
detailed investigation of additional material will be necessary
for a better understanding of these features. So far as is known,
all the species investigated here appear to be normal, sexual
species.

Cytological examination of preserved buds of Erythroxyvlum
havanense Jacq.(Plowman & Davis 3563), E. Ulei O.E. Schulz
(Plowman 6189) and E. areolatum L. (Kress s.n.) proved un-
rewarding. The smallest buds (ca. 0.5 mm. diameter) showed
pollen already formed. Furthermore, preserved material of
these wild species was difficult to work with because of tissue
hardness, preventing good squash-preparations. In the future,
better results may be obtained by special pretreatment of buds
or by examining root tips when material is available.

204

It is clear that much remains to be done in the cytology of
Erythroxylum. The study of cytogenetics has been essential in
determining the origin and evolution of numerous cultivated
plants. Erythroxylum should be no exception. Of initial, pri-
mary importance is an investigation of karyotypes and chro-
mosomal behavior in the main cultivated forms of E. coca and
E. novogranatense, as well as in related wild species.

LITERATURE CITED

Darlington, C.D. and E.K. Janaki Ammal. 1945. Chromosome Atlas of
Cultivated Plants. Allen and Unwin. London.

Heitz, E. 1929. Heterochromatin, Chromocentren, Chromomeren. Ber.
Deutsch. Bot. Ges. 47: 274-284.

Mangenot, S. and G. Mangenot. 1958. Deuxieme liste de nombres chromo-
somiques nouveaux chez diverses Dicotyledones et Monocotyledones
d'Afrique occidentale. Bull. Jard. Bot. Bruxelles 28: 315-329.

Mangenot, S. andG. Mangenot. 1962. Enquete sur les nombres chromosom-
iques dans une collection d’especes tropicales. Revue Cytol. Biol. Veg.
25: 411-447.

Mehra, P.N. and P.K. Khosla. 1969. /n A. Love, IOPB Chromosome Num-
ber Reports XX. Taxon 18: 215.

Tijo, J.A. and A. Levan. 1950. The use of oxyquinoline in chromosome
analysis. An. Estac. Exp. Aula Dei 2: 21-64.

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WO TAXOUHRLAYY NI SHAHN ON AINOSOWOUHD MAN (OME ATAVL

207

Plate 17.

EXPLANATION OF PLATE

Fig. 1-6. 1. Microsporocyte of Erythroxylum Coca, Plowman 6183.

X 1000.

2. Microsporocyte of Ervthroxvlum Coca, Plowman 6165. X 1000.
3. Microsporocyte of Ervthroxvlum novogranatense, Plowman
3685. X 1300.

4. Microsporocyte of Erythroxylum novogranatense, Plowman
6180. X 1300.

5. Root tip cell of Ervthroxvlum novogranatense, Plowman 6275. X
2800.

6. Root tip cell of Ervthroxvlum novogranatense var. truxillense,
Plowman 6250. X 2900.

208

209

BOTANICAL MUSEUM LEAFLETS

SOMA BROUGHT UP-TO-DATE*

R. GORDON WASSON

Already a decade has passed since the publication of
the English edition of our SOMA: Divine Mushroom of
Immortality. SOMA first made its bow to the world in an
expensive de luxe format. That edition is exhausted and
copies are quoted in the auction markets for rare books at
a considerable advance in price. The hard cover edition is
also exhausted. Of the paperback edition there has beena
succession of printings. It sells steadily, especially in
university centers of the English-speaking world.

When I undertook to study the Soma enigma, I ap-
proached it from outside the Sanskrit discipline, from
ethnobotany. After all, if a plant is to be identified, why
not turn to botany? And if the plant, when its juice was
ingested, inspired its devotees with glowing rapture and
adoration, why not have recourse to someone conversant
with plant hallucinogens? The West has known the
RgVeda for more than a century. The Vedists have failed
to identify the Soma plant, if indeed they ever made any
effort to do so: of such effort I have found no serious
trace, however strange this may seem to non-Vedists.

Naturally I needed a Vedist to work with me. My first
stroke of good fortune was to win the enthusiastic cooper-
ation of a young and brilliant Sanskrit scholar, Wendy
Doniger O’ Flaherty, who holds doctorates in Sanskrit and
Indian Studies from Harvard and Oxford Universities and

*Also will appear in the Journal of the American Oriental Society 99: 1979.
Published monthly except during July and ‘August by the Botanical Museum, Harvard University,
Cambridge, Massachusetts 02138. Printed by Harvard University Printing Office. Subscription:
$25.00 a year, net, postpaid. Orders should be directed to Secretary of Publications at the above
address. Second-Class Postage Paid at Boston, Mass.

who has lately become Professor of the History of Reli-
gions and Indian Studies at the University of Chicago. She
is the author of Part II of our SOMA, ‘The Post-Vedic
History of the Soma Plant’.

My method was to study Louis Renou’s translation of
the RgVeda, as far as he had gone before his death, espe-
cially but by no means exclusively Mandala IX, as well as
all his essays and commentary on the Vedic text. I also
found richly rewarding (1) Abel Bergaigne’s three vol-
umes on La Religion Védique, packed full as they were
with brilliant apercus, even prophetic in some instances,
although he never suspected that a mushroom could be
the answer to the identity of Soma, and (2) L’Agnistoma
of W. Caland and V. Henry, in which they give a slow-
motion picture, as it were, of the Soma sacrifice. There
were giants in the land in those early days of Vedic
studies. My transatlantic correspondence with Dr.
O'Flaherty was voluminous, as we constantly exchanged
ideas and I raised questions with her.

Early in my inquires it seemed to me that a mushroom
was the answer to many passages in the Vedic text and
was contradicted by none. I went over the ground again
and again and explored further and further. The result was
that | accumulated many texts, each supporting my mush-
room surmise, and I found none contradicting it. Not one
of my passages could be considered proof — far from it —
but the accumulation of many such passages would carry
increasing conviction. Here is the technique of overlap-
ping circles and ellipses, which is a commonplace in the
natural sciences. To this day Darwin's theory of the Ori-
gin of Species has not been conclusively proved, but the
accumulation of evidence in its favor has been over-
whelming and convincing; no one of consequence chal-
lenges it. When we published our SOMA I waited with
bated breath for fear a text from the RgVeda would be
produced incompatible with our mushroom hypothesis:
none has been offered. Instead, some Vedic and Sanskrit
scholars have produced most helpful, even startling, sup-
port.

The beauty of our case is that our evidence is linked in
parallel and not in series, and one or another of our separate
arguments can be shot down and yet our overall conclusions
will hold good.

Of primary interest to the readers of the French edition is the
public reception that the book has had from leading scholars in
the Sanskrit world, especially those competent in the RgVeda.
This reception has been conspicuously marked by silence: so
far as I know, there have been only three full-dress reviews by
leading Vedic scholars, two unfavorable and one favorable. To
each of the unfavorable reviews I have published a detailed
reply.

But that silence extends only to the printed word. The dis-
cussions among Sanskritists and especially Vedic scholars
were for a while lively. Beyond the confines of those limited
disciplines, among botanists, anthropologists, indologists,
sinologues, students of religion, etc., etc., the references to our
mushroom surmise in lectures, discussions, and publications
have been steady and, so far as I have knowledge, all lauda-
tory. But these people are not Vedists.

For generations the Vedic and Sanskrit fraternities have
been an ingrown community: the outside world of scholarship
and science has largely left them to themselves to till their rich
gardens in peace, and in turn they have largely ignored that
world. I hear tell of fratricidal infighting in the Sanskrit family,
but however that may be, they are unaccustomed to invasion
from the outside and many do not welcome intruders: they
close their ranks when the watchman cries ‘Stranger!’ In Vedic
time the Brahmans were the exclusive custodians of the Se-
crets of Soma: today some of the Vedic scholars occupying
exalted chairs seem to have replaced the Brahmans in their
attitude of superior exclusivity, though of course these Vedic
scholars are not possessed of the Secrets.

In SOMA a Vedist and ethnomycologist, working as a team,
think we may have found the answer to the Soma enigma. As I
shall show shortly, invaluable support for our position has been
forthcoming from those scholars whose minds are open to new
ideas.

There are problems in the humane letters as well as science

213

that do not lend themselves to final answers. In our SOMA we
advanced with all modesty a thesis pointing to an identification
of the Soma plant. We marshalled evidence favoring (not prov-
ing) this identification and hoped that the evidence was enough
to interest specialists and to encourage them to look further.

When Darwin announced his theory of evolution, there were
those Biblical scholars who took the position that if his theories
challenged the literal truth of the Bible he must be wrong and
the Bible was of course (so they said) right. External evidence
on the Bible has revolutionized conceptions of it in the past
century and the RgVeda is no more immune to detached study
than the Bible. As compared with the theory of evolution, the
controversy over Soma in the RgVeda is the same en petit, en
tres petit. There is a category of Indo-Iranian exegete who lives
immured in his enclave, unfamiliar with the outside world.
Fortunately not all Indo-Iranian scholars are of that breed. We
must remember, however, that exegetes, useful — indeed in-
dispensable — as they are, are often at the end of the line when
it comes to assessing great poetry.

Some of the most rewarding years of my life I have spent
under the spell of the line of seminal Vedic and Sanskrit schol-
ars whom I have already mentioned: Bergaigne, and Caland
and Henry, and Renou, as well as many other figures of compa-
rable stature in the English-speaking world. That succession of
learned and intuitive scholars seems, at least momentarily, to
have fallen off somewhat, with Renou’s tragic death and the
death of the R.P. Jean de Menasce, O.P. My conversations
with Renou in his Paris home and his country place in Norm-
andy, the letter that Menasce wrote me from his death-bed,
make me certain that had they lived, whether they agreed or
disagreed with me, the breadth of their understanding would
have been inspiring, their pronouncements discriminating and
helpful. They would have grasped the possibilities that lay in
interdisciplinary exchanges — long overdue — for Vedic
studies. No one could ever say that they possessed constipated
mentalities.

214

FRESH EVIDENCE
A. THE PUTKA OF THE SANTAL

At an early stage of our mycological inquiries the late Georg
Morgenstierne, the distinguished Norwegian scholar specializ-
ing in the babel of tongues spoken in Nuristan, drew my atten-
tion to an oddity of the language spoken by the Santal, a tribe
numbering some millions living in scattered villages of Orissa
and Bihar in India. Their language is neither Indo-European
nor Dravidian: in India it heads a third much smaller family
called Munda. A trait of these languages is that they possess no
genders: Munda speakers know not our masculine, feminine,
neuter categories. For them all creatures and objects are either
animate, possessing a soul, or inanimate, devoid of soul. The
division between the two seems to us somewhat arbitrary,
enshrining conceptions that prevailed long long ago. Thus the
sun, moon, stars were and are conceived of as animate. On the
other hand the whole of the vegetable kingdom is inanimate —
with one exception, a single species of mushroom* called
‘putka’. What was this mushroom and why was it animate?

In 1965 I visited the Santal Parganas (as they are called) in
Bihar, in and around Dumka. I questioned as many of the older
people as I could. Mrs. A.E. Stronstad, the wife of the Norwe-
gian missionary, most graciously was serving as my interpre-
ter. | found the Santal an endearing people, gentle, eager to
help, obviously candid. None of them that I spoke with knew
either English or Hindi. The Santal are, unlike the Hindus,
mycophiles. My best informant was an elderly woman, Ludgi
Marndi. She told me there was one mushroom that caused
inebriation. Was it the putka? No, definitely not. What was the
putka? We were in January and none were around at that time
of year. Why was it animate? No one knew: apparently the
reason had been lost in the depths of the past. A few informants
pointed out hesitatingly that putka abounded in the sacred
grove of sarjom trees growing near every village. (Santal sar-
jom = Hindi sal = Shorea robusta) Was this perhaps the

**Mushroom in French is champignon, but unlike ‘mushroom’ in English, champig-
non embraces the whole fungal world even to the microscopic species. As this paper
was to be translated into French, ‘mushroom’ in it embraces ascomycetes as well as
basidiomycetes.

215

reason? But the sarjom trees themselves were not animate, and
putka grew abundantly elsewhere: how then could that be the
reason? No answer.

Just as I was about the leave the Santal country I engaged
Ludgi Marndi in one final talk. We went over the same ground.
Suddenly she leaned forward and in a whisper made a most
curious remark, which she said was her guess as to why the
putka were animate: the putka must be eaten on the very day
they are gathered, ‘for within twenty-four hours they will stink
like a cadaver’. She was whispering. She was so earnest that I
wrote down in my journal at once what she had said. But I
confess that it meant nothing to me.

In 1967 the mycologist Roger Heim, then directeur of the
Museum National d’ Histoire Naturelle and former President of
the Académie des Sciences, and I, he from Paris and I from
New York, journeyed to the Santal villages of Orissa and
Bihar. We went in July. We found putka growing in abundance
and Heim identified them. They were fungi without inebriating
potency. The Santal gathered these hypogenous fungi just as
they were appearing on the surface, little dark-brown globular
mushrooms. We questioned villagers along our whole route but
no one could tell us why these mushrooms were animate. When
we published our paper on our trip in Les Cahiers du Pacifique
#14, in September 1970, we offered that baffling sentence of
Ludgi’s to our readers, though it still meant nothing to us.

It meant nothing to us until the April-June 1975 issue of the
Journal of the American Oriental Society (JAOS) came out.
Then it developed that her remark meant everything to us. That
JAOS paper explained 1) Ludgi’s baffling remark, 2) why the
putka were animate, 3) and the etymology of putka. It fortified
even further, some say immeasurably, the thesis that we had
presented in SOMA: Divine Mushroom of Immortality.

The paper in the JAOS was written by Stella Kramrisch. She
has spent 27 years in India as Professor of Indian Art at the
University of Calcutta. She also taught concurrently at the
Courtauld Institute, London University. From there she had
gone to the University of Pennsylvania as Professor of South
Asian Art. She was and is now the Curator of Indian and
Himalayan Art in the Philadelphia Museum of Art and Profes-

216

sor of Indian Art at the Institute of Fine Arts of New York
University.

P.O. Bodding in the preface to his five volume Santal Dictio-
nary (Oslo, 1929-1936, I:xiv) had drawn attention to a note-
worthy fact:

Strangely enough, the Santals use some pure Sanskrit words,
which, so far as I know, are not heard in present day Hindi.

He might have added that, at least in the instance we are
considering, the word has disappeared fromal/ Sanskrit verna-
culars. Professor Kramrisch wins a rich accolade for discover-
ing that Santal putka was a loan word from Sanskrit, putika, a
plant heretofore unidentified. She had read our paper in the
Cahiers du Pacifique, and saw immediately that putka was
simply the Sanskrit putika. It is cognate with our word ‘putrid’,
and the stench of the putka lives up to its Sanskrit name. The
putika was mixed with the clay that went into the ceremonial
making of the Mahavira pot. Its offensiveness was turned into
fragrance when the pot, held by tongs, was fired in the course
of the rite. It was the earliest surrogate for Soma. No one had
ever known what plant it was. Not even Bodding had hit on its
identity. In our SOMA Dr. O'Flaherty had mentioned putika
four times but I failed to link it with the putka: Stella Kramrisch
in the JAOS had identified it with finality. Professor Kramrisch
wrote:

The identification of Putika, the Soma surrogate, supplies
strong evidence that Soma indeed was a mushroom. Putika
integrated into the Mahavira pot played its part in the mystery of
the Pravargya sacrifice. That putka-mushrooms should be
known, to this day, as ‘endowed with a soul’ witnesses amongst
the Santal of Eastern India a memory of the numinous emanating
from the indigenous Indian Soma substitute... . (Vol. 95:2,
p. 230 col. 2)

A gap of more than two and a half millennia, a transfer from
Aryan priestly symbolism to tribal belief, the tribe adopting a
Sanskrit name with but little change into its own language, the
survival of this name in a Munda language, in a region at a
considerable distance to the east from the ancient center of
Brahmanical sacrifices, all this did not impair the ongoing myth
of Putika. This species is known to the Santal as ‘endowed witha
soul’. It is distinct from other mushrooms, from all the vegetable
kingdom, as being numinous. The odor of sanctity clings to this

217

mushroom, however pejorative its telling name... . (p. 233
col. 2)

The mantle of King Soma had fallen on this notable mush-
room, which inherits the glory of Soma for whom it is a surro-
gate. Its heyday is in the Pravargya ritual. Then the mushroom is
lost to us in a millennial darkness from which, miraculously, it
emerges ‘endowed with a soul’, amongst the aboriginal Santal of
Eastern India in our own day. . . . (p. 235 col. 2)

How amazing it is that through a concatenation of happen-
stances we are able, here in 1978, to define the precise species
of the plant putika, which the Aryans, perhaps 3000 years ago,
adopted as a surrogate from Soma! Preserved as in a time
capsule in the Santal country and called in the Santal language
putka, for 3000 years it waited to be discovered and its meaning
made known by Roger Heim, Directeur of the Museum Na-
tional d’ Histoire Naturelle, by the Austrian-born art historian,
Stella Kramrisch, and by the ethnomycologist Gordon Was-
son, two of us working as a team, Heim and Wasson seeking
the reason why putka, alone in the vegetable kingdom, was
animate and not finding the answer, not until Kramrisch re-
vealed it to them, on the strength of evidence we had gathered
but had failed to understand! According to the laws of Manu,
mushrooms have been forbidden for thousands of years to
Hindus of the twice-born castes. This makes it all the more
significant that a mushroom was at one time the Holy of Holies.

The late R.C. Zaehner, a distinguished scholar in the Indo-
Iranian field and a professor in Oxford, in his review of our
SOMA in The Times Literary Supplement, 22-5-1969, gave as
the main argument against my mushroom theory the fact that
when Soma came to be abandoned other mushrooms were
never the substitutes. He did not live long enough to learn that
the primary surrogate, the putika, was a mushroom.

Manfred Mayrhofer in his Concise Etymological Sanskrit
Dictionary presented two identical words that he spelled
puatikah: 1) ‘foul, stinking’, and 2) ‘a species of plant serving as
a substitute for the Soma plant’. Fortunately, in his ** Additions
and Corrections’ at the end of Vol 3, he was able to cite
Professor Kramrisch’s paper and thus reconcile and unite in
one word what he had presented as homonyms.

218

B. THE COLOR OF SOMA.

We said in SOMA that hari in the RgVeda meant red when
that color word was applied to Soma, which we think was A.
muscaria, amushroom that throughout Eurasia, when mature,
is almost always an intense and striking red. John Brough says:
‘I have been unable to find any evidence that any shade of red is
included in the colour-range denoted by hari’, and winds up his
discussion by declaring that ‘red is absolutely excluded’.

Scholars know that to arrive at the value of color words in
texts 3000 and more years old calls for intense linguistic re-
search, citation by citation, pinpointing as close as possible the
time and place of each citation, and the study of other overlap-
ping color words, and of cognate color words in surrounding
languages, efc., etc. In Brough’s discussion of hari there is
none of this, only his ipse dixit.

Sir Harold W. Bailey, Brough’s predecessor in the chair in
Cambridge that Brough now holds, undertook a study of the
meaning of ZAR, cognate in Khotan Saka to Vedic hari. His
findings appeared in a collection of learned papers: Mémorial:
Jean de Menasce, published in 1974 as #185 by the Fondation
Culturelle Iranienne. I will quote only one sentence of his
conclusions:

Important for the Irano-Indian period is the corresponding Old

Indian vocabulary. Here hari-, harit-, harita- has the same wide
range from red through orange to yellow and green. (p. 372)

Bailey mentions neither Brough nor me.

C. THE ETYMOLOGY OF *“SOMA’

‘Soma’ has been known in the West for almost two centuries
and has been a focus of scholarly attention for most of that
time. Yet Indic scholars including Brough have docilely ac-
cepted the ‘etymology’ that the Brahmans have given it and say
that so- is the Vedic su-, meaning the ‘pressed thing’, from the
liturgical act in the Soma sacrifice when the plants were noisily
pounded with stones on hide-covered boards. As Bailey said in
1971 in Tokyo, this is ‘a poor kind of way to designate a
sacrificial plant of great potency’. Indeed, the plant must have

219

had a name before the liturgy was devised, but then we are to
suppose that that name was utterly lost, and the plant came to
have no name at all! Is this conceivable? But in the Brahman
world etymologies were creatures of word-play, puns, poetic
fancy.

Bailey has suggested an alternative etymology, which he has
presented briefly in three publications, listed below. Instead of
so-ma-, he would break the word thus: som-a- and then it
would be linked with the Indo-European root for ‘fungus’, of
which German Schwamm, Latin fungus, Greek spongia are
just afew examples in Europe. Many Vedic and Sanskrit schol-
ars display a low flash-point when this fungal theme 1s
broached. Why this ts so is baffling. It has been suggested to me
that a spiritual osmosis from the Brahmans is the explanation,
for converts are proverbially more Papist than the Pope. |
prefer to think that as I was the first to suggest a mushroom,
they may resent the irruption of an outsider into their domain.
Let them be cautious: Bailey's suggestions have a way of
turning out right in the end. After all, the Law of Parsimony has
its place here: as between two unproved etymologies, the
plausibie one has an immense advantage over the one that
strains credibility.

Bailey presented this etymology orally at Canberra, at the
28th International Congress of Orientalists in January 1971. He
next developed the theme in the Memoirs of the Research
Department of Toyo Bunko (The Oriental Library), No. 29,
Tokyo, in 1971, pp 8 and 1S. He discussed it again in his paper,
‘A Half-Century of Irano-Indian Studies’, which he read as an
informal lecture after he was awarded the Triennial Gold Medal
of the Royal Asiatic Society on 13 April 1972 and which was
published in the Society's Journal No 2, 1972, p 105. He
returned to the theme in his contribution to Mithraic Studies,
edited by John R. Hinnells and published by Manchester Uni-
versity Press in 1975, p 19, ftnt 38.

D. AJA EKAPAD

Through oversight I failed to include in our SOMA a discus-
sion of Aja Ekapad, the designation of a deity mentioned six

220

times in the RgVeda, always in hymns to the Vi’svadevas.
This name means, as Abel Bergaigne pointed out and everyone
agrees, non-né unipéde ‘not-born singlefoot’. There has been
much scholarship expended on this name for a god: the ‘Not-
born Singlefoot’, but no one has seen how perfectly it fits a
mushroom. A mushroom is uwnipéde: to this day every German
child still learns at his mother’s knee the riddle:

Sag’ wer mag das Mannlein sein

Das da steht auf einem Bein?

Children: Gluckpilz! Fliegenpilz!

Less familiar to us urbanized westerners, living divorced
the land and unfamiliar with the elementary facts of nature that
our ancestors well knew, isnon-né. All plants are born of seed,
except mushrooms. They are non-né. The spores of a mush-
room are too small for the eye to see: they require a micro-
scope. Mushrooms were, as Early Man saw it, miraculously
conceived, by the lightningbolt of the Almighty in mother earth
softened by the rain. There is a verse in the RgVeda that says
Parjanya, the god of Thunder, was the father of Soma, and
according to another verse, ‘the gods, those fathers with a
commanding glance, laid the Somic germ’. In Aja Ekapdd we
have the perfect binomial for a mushroom!

Aja Ekapad is accompanied in five citations by another
divinity, Ahi Budhnya, the serpent of the depths, a chthonic
being who invariably guards the holy plant throughout Asia.
That Aja Ekapdad is another name for Soma, perhaps already
archaic in Vedic times, finds support in RgVeda X 65.13, where
the epithet proper to Soma, * Mainstay of the Sky’, is applied to
him. In RgVeda X 82.6 Aja appears, not now linked
to Ekapad, but to the navel, nabni, also endlessly applied to
Soma, and thus two seemingly disparate metaphors are recon-
ciled in the divine mushroom, both proper to Soma.

Here is Renou’s translation of the second half of the verse we
are discussing, X 82.6:

Dans le nombril du Non-ne (est) fixe le Un (comme les rais sur la
roue), en lequel tous les etres se tiennent-depuis-toujours.

In the navel of the Not-born (is) fixed the One (like the spokes on
the wheel), in which all beings stand from all time.

22)

He has interpolated — clearly indicated by the parentheses —
‘like the spokes on the wheels. Where did Renou get these
words or the idea that they contain? Obviously he did not
invent this gloss. Dr. O’ Flaherty has sought the source in the
obvious places and has not found it. Perhaps it occurs in the
other Vedas or the Brahmanas. It might prove of vital interest,
for the gills of a mushroom make one think of the spokes of a
wheel: attached to the stipe as to the navel.

I will end this esssay on a lighter note.

One aspect of our SOMA that was utterly repugnant even to
my friends and supporters in Europe and America was my
discussion of Soma-urine. Without exception they were hor-
rified by the thought of drinking urine, and filled with disbelief.
This surprised me as I had thought we in this generation had
overcome our parochial squeamishness. In my Rejoinder to
Brough I handled this topic with special care, citing contem-
porary evidence to indicate that urine drinking goes on to this
day in India and I think in most of the world’s population. Early
Man, probably everywhere, recognized that in urine he pos-
sessed an aseptic fluid useful for wounds, serving also in reli-
gious observances, and perhaps wholesome for ingestion.

In the fall of 1977 Morarji Desai, Prime Minister of India,
made known to the world that he drinks his own urine every
day, for his health’s sake. Little did the Prime Minister know
how this innocent announcement would affect the post and the
telephone calls of Gordon Wasson in Danbury, Connecticut.
All sorts of people, friends and even strangers, got in touch
with me to congratulate me. One Sanskrit scholar, a friend,
said that now the last barrier to my Soma theory was demo-
lished! Another man, engaged in the public relations business,
a stranger who scarcely knew me, asked me how I had con-
trived such an interview. What a coup! Did I have a personal
contact with the Prime Minister, or know someone who knew
him? Alas, I could claim no credit.

Pps

WASSON’S SOMA: IMPORTANT REVIEWS
AND PERTINENT COMMENTARY

Bailey, H.W.: (1) ‘The Range of the Colour ZAR in Khotan Saka Texts’,
Mémorial: Jean de Menasce, Fondation Culturelle Iranienne,
Louvain, 1974; pp. 369-374: our quotation is from p. 372.
(2) ‘Trends in Iranian Studies’, Memoirs of the Research
Department of the Toyo Bunko (The Oriental Library) No. 29,
Tokyo, The Toyo Bunko; see pp. 8 and 15.

(3) ‘A Half-Century of Irano-Indian Studies’, Journal of the
Royal Asiatic Society, 1972:2, pp. 99-110; see p. 105.

(4) ‘The second stratum of the Indo-Iranian gods’, Mithraic
Studies, edited by John R. Hinnells, Manchester University
Press, 1975; see ftnt. 38.

Brough, John: ‘Soma and Amanita Muscaria’, BSOAS, 34:2, 331-362. Re-
view Article.

Gershevitch, Ilya: ‘An Iranianist’s View of the Soma Controversy’, Mémo-
rial: Jean de Menasce, Fondation Culturelle Iranienne, Lou-
vain, 1974; pp. 45-75.

Heim, Roger, and R.G. Wasson: ‘Les Putka des Santals Champignons doues
d’une ame.” Cahiers du Pacifique N° 14, Sept. 1970; pp. 60-85,
11 planches hors texte, 6 en couleur. The Ludgi Marndi cita-
tion is on p. 65.

Ingalls, Daniel H.H.: (1) The New York Times, Sunday Book Review Sec-
tion, 5 Sept. 1971.

(2) JAOS, 91:2, 1971.

Kramrisch, Stella: ‘The Mahavira Vessel and the Plant Putika’, JAOS 95:2,
April-June 1975, 222-235.

Kuiper, F.B.J.: Indo-Iranian Journal, 12:4, 1970, pp. 279-285. Review Arti-
cle.

La Barre, Weston: American Anthropologist, Vol. 72, 1970, 368-373. Re-
view Article.

Lévi-Strauss, Claude: ‘Les champignons dans la culture: A propos d'un
livrede M.R.-G. Wasson’, L’Homme, Vol. X, Cahier 1, 1970,
pp. 5-16. Review Article.

Schultes, Richard Evans: Journal of Psychedelic Drugs, Vol. 3:2, Spring
1973. Review.

Wasson, R.G. (1) ‘Soma: Comments Inspired by Professor Kuiper’s Re-

view’, Indo-Iranian Journal, 12:4, 1970, pp. 286-298. This
article appeared in the same issue immediately after the
Kuiper review.
(2) ‘Soma of the Rig Veda: what was it?’, JAOS, 91:2, April-
June 1971. This paper was republished later by the American
Oriental Society in their Essay Series, No. 7, along with
Daniel Ingalls’s review that had appeared with itintheJAOS.
(3) ‘Soma and the Fly-Agaric: Mr. Wasson’s Rejoinder to
Professor Brough’, with Introduction by Richard Evans
Schultes. Published by the Botanical Museum of Harvard
University, Cambridge, Mass. 1972.

a5

BOTANICAL MUSEUM LEAFLETS VoL. 26, No. 6
JUNE 30, 1978

DE PLANTIS TOXICARITS E MUNDO NOVO
TROPICALE COMMENTATIONES XXIII

ETHNOPHARMACOLOGICAL NOTES FROM
NORTHERN SOUTH AMERICA

RICHARD EVANS SCHULTES

Increasing evidence indicates that the indigenous population
of the northwesternmost part of the Amazon Valley possesses
an almost unsurpassed wealth of knowledge of biodynamic
plants. There are few parts of the world where native people
display such a basic understanding of plants of use as poisons,
narcotics or medicines.

Until recently, this corner of the great hylea has not been in
danger from penetration by outside civilization. How long this
happy state of affairs will continue is open to doubt.

In order to preserve some of this native folk lore, | have been
publishing a series of articles summarizing some of the ethno-
pharmacological information which my students and I have
collected in field work from 1941 to the present.

Most of the following notes pertain to the wealth of ethno-
pharmacological knowledge of Indians of the northwestern
Amazon — primarily in Colombian territory, but occasionally
from adjacent parts of Brazil, Ecuador and Peru.

Voucher specimens cited are preserved in several herbaria:
the Economic Herbarium of Oakes Ames and the Gray Her-
barium of Harvard University, the New York Botanical Gar-
den, the Herbario Nacional de Colombia (Bogota) and the
Instituto Nacional de Pesquisas da Amazonia (Manaos).

The families are arranged in accordance with the Engler and
Prantl system, and the genera are alphabetically enumerated
under their respective families.

LYCOPODIACEAE

Lycopodium cernuum Linnaeus, Sp.Pl. (1753)1103.
225

CoLomsiaA: Comisaria del Amazonas, Rio Amazonas, vicinity of Leticia.
August 29 - September 12, 1966. R.E. Schultes, R.F. Raffauf et
D. Soejarto 24038.

A Dragendorff spot-test for alkaloids was positive for this
common species of Lycopodium.

Alkaloids have recently been detected in about a dozen
species of Lycopodiaceae. Nicotine and quinolizidine bases
account for most of the lycopodiaceous alkaloids (Tyler, V.E.:
Lloydia 24(1961)58).

PIPERACEAE

Peperomia macrostachya (Va/il.) Dietrich var. nematostachya

(Link.) Trelease et Yuncker, Piperac. N. S. Am. 2(1950)661.

COLOMBIA: Comisaria del Vaupes, Rio Kananari, Cerro Isibukuri. ** Epi-
phyte’’. March 8, 1951. R.E. Schultes et 1. Cabrera 13260. —
Rio Vaupes, Mitu and vicinity. R.E. Schultes, R.F. Raffauf et
D. Soejarto 24178.

The leaves of this climbing epiphyte are boiled, and the
resulting tea is taken by elderly members of the Taiwano tribe
to relieve difficulty in urinating. It is said also to be an effective
febrifuge.

A Dragendorff spot test for alkaloids (Schultes, Raffauf et
Soejarto 24178) was negative.

Piper hostmannianum (Mig.) C. DeCandolle in DeCandolle
Prodr. 16, pt. 1(1869)287.
COLOMBIA: Comisaria del Amazonas, Rio Amazonas, vicinity of Leticia.

“Small bush. Leaves aromatic.”” August 29 - September 12,
1966. R.E. Schultes, R.F. Raffauf et D. Soejarto 24018.

In the Leticia region, a poultice of the crushed leaves is
applied to warts in the belief that this hastens their disappear-
ance. The plant is known as cordoncillo, a name applied to
many species of Piper in reference to the inflorescence (cor-
doncillo = *‘shoe string’’).

Piper interitum 7release ex Macbride in Field Mus. Publ. Bot.
18, No. 357(1936)176.

226

PERU: Departamento del Loreto, Zapote, Alto Rio Purus. “Tree about 10
meters high. Lowland forest’’. October 15, 1968. L.Rivier 21.

The dried leaves and roots are pulverized and used as a
substitute for tobacco snuff. The Kulina Indian name is frets.

Pothomorphe umbellata (L.) Miquel, Comm. Phyt. (1840)36.

CoLomBIA: Comisaria del Putumayo, Rio Guamues, San Antonio. **Secon-
dary growth. Herb 5-6 feet.”’ September 5, 1966. H.V. Pinkley
42].

The Kofan Indians prepare an arrow poison of this plant
alone (the bark of the lower stem and root is scraped and
boiled) or mixed with other plant ingredients, especially for
hunting monkeys and wild pigs. The Kofan name is a-nama-he
se-hé-pa.

This species has been employed as a strong diuretic in Brazil
(Peckholt, T.: Pharm. Rundschau 12(1894)240,285), but there
appears to be no chemical constituent which would make the
bark active as an ingredient of curare.

MyYRICACEAE

Myrica parvifolia Bentham, Pl. Hartw. (1846)251.

CoLomBiA: Departamento de Cundinamarca, Bogota, Cerro above La Cita.
May 10, 1946. R.E. Schultes 7111.

Bundles of the leaves and stems of this shrub are burned in
huts in the cool, humid highlands around Bogota for the aro-
matic smoke which is believed to relieve congestion casued by
the frequent pulmonary troubles of these people.

POL YGONACEAE

Rumex obtusifolius Linnaeus, Sp. Pl. (1753)335.

CoLoMBIA: Comisaria del Putumayo, Valle de Sibundoy, Sibundoy. Alt.

2225-2300 m. May 29, 1946. R.E. Schultes et M. Villarreal 7610.
— Same locality. ‘‘Lengua de vaca’’. February 20, 1963. P.J.
Chindoy B. 97. — Same locality. **Garden and waste places,

very frequent. One or two most persistent, most frequent weeds
in valley’’. April 1, 1963. M.L. Bristol 704. — Same locality.
“Lengua de vaca. Open pasture; infrequent."’ May 8, 1963.
M.L. Bristol 969.

227

The Kamsa Indians of Sibundoy employed the roots of Rumex
obtusifolius in decoction as a strong laxative.

The roots contain 1,8-dihydroxynaphthaline (Hesse, O.:
Ann. Chem. 305 (1896)291).

MENISPERMACEAE

Curarea tecunarum Barneby et Krukoff in Mem. N.Y. Bot.

Gard. 22(2)1971)12.

BRAZIL: Estado do Amazonas, Rio Cunhua, Deni Indian village. November

28, 1971. G.T. Prance, R.J.M. Maas, D. Woolcott et al. 16453.

CoLoMBIA: Comisaria del Putumayo, Rio Guamiles, Salvador. ‘‘ Arrow
poison” August - September 1963. C. Naranjo et G. Wiederhold
16. — Comisaria del Vaupes, Rio Macaya, vicinity of Cachivera
del Diablo. ‘‘Said to have been used formerly by Karijona In-
dians in arrow poisons. Vine. Fruits yellow, very bitter.’’ May
1943. R.E. Schultes 5526. — Rio Macu-Parana, lowland forest.
‘Root scrapings are used in preparing arrow poison mixture by
Bara-Maku. Awa-puh’ (species of monkey root). June 1-8, 1970.
P. Silverwood - Cope 23.

Curarea tecunarum is well recognized as one of the principal
sources of an especially strong curare in the northwestern
Amazon. Its use as a contraceptive, however, is not widely
known. According to Prance, Maas, Woolcott et al. 16453, the
stem is crushed and placed in water, stirred and taken as a

contraceptive by the Deni Indians.

Telitoxicum peruvianum Moldenke in Brittonia 3(1938)45.

CoLomBIA: Comisaria del Vaupes, Rio Piraparana, Cano Teemeena, (Lobo
Igarape). ‘‘Small tree. Fruit dark green. Barasana: bo-dé-mee-
see’. September 10, 1952. R.E. Schultes et 1. Cabrera 17340.
The Makuna Indians value Telitoxicum peruvianum as an
important ingredient of the curare that formerly was prepared
by medicine men of the Rio Piraparana. The Barasana believe
that application of crushed leaves to ulcers and similar infec-
tions aids in cleansing the wounds.

ANNONACEAE

Anaxagorea sp.

EcuADorR: Napo, Rio Aguarico, Dureno. **Tree.’” December 12, 1955. H.V.

228

Pinkley 16. — Same locality. ‘‘Small tree, 6-8 feet.”’ October 19,
1966. H.V. Pinkley 522.

The bark of the root of this treelet is an ingredient of curare
amongst the Kofans. The Kofan name is ko-yo-vi-fd-nti.

Cyanogenesis is reported from a Philippine species of
Anaxagorea (Hegnauer, R.: Chemotaxonomie der Pflanzen
3( 1964) 121)

Guatteria Duckeana R.E. Fries in Acta Horti Berg. 12 (1939)
468.
COLOMBIA: Comisaria del Vaupes, Rio Vaupes, Cachivera de Tatu. **Tree

45 feet. Flowers green.’ October 20, 1966. R.E. Schultes, R.F.
Raffauf et D. Soejarto 24377.

All parts of this plant tested very positive for alkaloids with
Dragendorff reagent.

Guatteria dura R.E. Fries in Acta Horti Berg. 12(1939)499.

CoLoMBIA: Comisaria del Vaupes, Rio Kuduyari. ‘Tree 18 feet. Flowers
yellow-green, cauliflorous.’’ October 10, 1966. R.E. Schultes,
R.F. Raffauf et D. Soejarto 24391.

The bark tested highly positive for alkaloids, the leaves
slightly positive, with Dragendorff reagent.

Unonopsis veneficiorum (Mart.) R.E. Fries in Acta Horti Berg.
12(1937)238.

COLOMBIA: Comisaria del Vaupes, right tributary of Rio Macu-parana. June
1-8, 1970. P. Silverwood-Cope 11.

This plant is reported to be an ingredient of a contraceptive
formula of the Bora-Maku who inhabit the region of the Rio
Piraparana in the Colombian Vaupes. The name of the medici-
nal preparation in Maku is we-wit-kat-ku (‘‘no children
medicine’’).

Unonopsis veneficiorum has long been known as an ingre-
dient of a type of curare in the northwestern Amazon. The plant
is alkaloidal (Hegnauer, R.: Chemotaxonomie der Pflanzen
3(1964)118).

Xylopia amazonica R.E. Fries in Acta Horti Berg. 12(1939)562.
229

COLOMBIA: Comisaria del Amazonas. Rio Apaporis, Soratama. **Large
tree. Flowers white. High land.’’ September 28, 1951. R.E.
Schultes et 1. Cabrera 14146.

Xylopia amazonica is valued by the Indians of the Rio
Apaporis in the form of a tea to induce sleep. The leaves and
stems are employed. Alkaloids, polyphenols, and essential oils
have been reported from Xy/opia (Hegnauer, R.: Chemotaxo-
nomie der Pflanzen 3(1964)118, 120).

Xylopia Benthamii R.E. Fries in Kgl. Sv. Vet.-Akad. Hand.
34, No. 5(1900)35.
COLOMBIA: Comisaria del Vaupes, Rio Kananari, Cerro Isibukuri. Alt. 2500

feet. ““Small tree. Pods white and red. Forest on slope near
summit. December 4, 1951. R.E. Schultes etl. Cabrera 14751.

According to the Taiwanos of the Rio Kananar1, a tea of this
plant is administered as a tranquilizer to people who have
experienced a great fright.

MyYRISTICACEAE

Osteophloem platyspermum (Poepp.) Warburg in Nova Acta

Acad. C. L. C. G. Nat. Cur. 68(1897) 162.

BRAZIL: Estado do Amazonas, basin of Rio Negro, Rio Uniciuxi, Maku
Indian village 300 km. above mouth. **Forest on terra firma. Tree.
25m. x 40cm. diameter. Flowers green. Sap drunk by Maku as cure
for coughs and colds. Maku = tugnebanpe. October 23, 1971.
G.T. Prance, R.J.M. Maas, D.B. Woolcott et al. 15571. — Rio
Negro, vicinity of Manaos, Reserva Ducke. April 11 - 14, 1972. R.E.
Schultes et W. Rodrigues 26126a.

The Maku drink the sap as a “‘cure for coughs and colds.””
Labourers in the Reserva Ducke near Manaos burn the leaves
and inhale the smoke to relieve asthmatic conditions.

Chemical studies of Osteophloem apparently have not been
effected.

LEGUMINOSAE

Acosmium nitens (Voe.) Yakovlev in Notes Roy. Bot. Gard.
Edinb. 29(1969)353.

230

BRAZIL: Estado do Amazonas, Rio Negro, Tapurucuara. ** Flowers white.
Small tree. Bark very alkaloid-positive; petiole and leaf negative.”
July 5- August 12, 1967. R.E. Schultes 24550 (Alpha-Helix Amazon
Expedition 1967).

The bark of this tree was indicated as an ingredient of curare
made in former times by the Tukano Indians.

Heterostemon mimosoides Desfontaines in Mem. Mus. Paris

4(1818)248.

CoLomBIA: Comisaria del Vaupés, Rio Vaupés, Mitu and vicinity. ‘Small
tree or bush. Flowers purple. Alkaloid-negative.’’ September 27

- October 20, 1966. R.E. Schultes, R.F. Raffauf et D. Soejarto
24171.

The flowers are said formerly to have been pulverized and
used for flavouring chicha.

Ox ALIDACEAE

Oxalis lotoides Humboldt, Bonpland et Kunth, Nov. Gen. et
Sp. 5(1821)187.

CoLtomsiaA: Comisaria del Putumayo, Valle de Sibundoy, Sibundoy. Alt.
2225-2399 m. May 29, 1946. R.E. Schultes et M. Villarreal 7607.

The Kamsa Indians prepare a tea of this plant which is taken
as a gargle to relieve chest and throat pains.

It is perhaps significant that the leaves of another species of
this genus, the Amazonian Oxalis Martiana Zucc., are pre-
pared in the form of a gargle to relieve pains of angina (Le-
Cointe: A Amazonia Brasileira 3(1943)108).

Leucoanthocyanines have been reported from some species
of Oxalis (Bate-Smith, E.C.: Journ. Linn. Soc. London
(Botany) 58(1962)95-173).

VOCHYSIACEAE

Qualea acuminata Spruce ex Warming in Martius, FI. Bras. 13,
pt. 2(1882)40.

CoLomBia: Comisaria del Vaupés, Rio Vaupés, Mitu and vicinity. **Small
tree. Flowers white and pink.’’ September 27 - October 20, 1966.
R.E. Schultes, R.F. Raffauf et D. Soejarto 24177.

2351

This plant tests alkaloid-negative with Dragendorff reagent.
A tea of the bark is valued as a taenifuge in the Mitt area.
The chemistry of Qualea is unknown.

EUPHORBIACEAE

Croton glabellus Linnaeus, Syst. Ed. X(1758)1275.

CoLomBIA: Comisaria del Amazonas, near mouth of Rio Loretoyacu and
Puerto Narino. “‘Fruit green. Small tree.’’ September 13-15,
1966. R.E. Schultes, R.F. Raffauf et D. Soejarto 24149,

Witoto Indians living in the vicinity of Leticia crush leaves of
this tree to poultice infected cuts and sores.

Croton glabellus has been reported to be alkaloid-positive
(Haynes, L.J. et K.L. Stuart: Journ. Chem. Soc. 1963 (1963)
1784, 1789).

Croton palanostigma K/otzsch in Hooker Lond. Journ. Bot. 2
(1843) 48.

BRAZIL: Estado do Amazonas, vicinity of Manaos, Reserva Ducke. **Small
tree. Flowers white’. April 11 - 14, 1972. R.E. Schultes et W.
Rodrigues 26124 1a.

The sap of this tree is applied to ulcers and boils to reduce
pain.

Mabea nitida Spruce ex Bentham in Hooker Kew Journ. 6
(1854) 367.

CoLoMBIA: Comisaria del Vaupes, Rio Vaupés, Mitu and vicinity. ‘*Tree 20
feet. Fruit rusty. Alkaloid-negative.’’ September 27 - October
20, 1966. R.E. Schultes, R.F. Raffauf et D. Soejarto 24170.

The oil extracted from the seeds was, according to Kubeo
informants, formerly rubbed into the scalp to prevent or delay
loss of hair.

BOMBACACEAE

Bombax globosum Aub/et, Pl. Guian. Fr. 2 (1775) 701.

COLOMBIA: Comisaria del Vaupes, Rio Kananari, Cerro Isibukuri. ‘Large
tree, 60 - 70 feet tall. Flowers yellow-white, petals yellow to-

232

wards tip.’ September 29, 1951. R.E. Schultes et I. Cabrera
14700.

The Taiwano Indians, who know this tree as ka-ne-wee-re,
gathered the ‘‘wool’’ from the ripened fruits for use, mixed
with latex or resin from a number of plants, to apply to cuts,
open sores or ulcers as a kind of protection during healing.

STERCULIACEAE

Herrania Camargoana R.E. Schultes in Bot. Mus. Leafl., Har-

vard Univ. 14(1950)120.

BRAZIL: Estado do Amazonas, Rio Cauaburi, Maturaca. **Single slender
trunk. Height 20 ft. In flood forest. Fruit brownish red with fleshy
pseudospines at junction of ribs and cross ridges. Alkaloid-nega-
tive.’ July 5- August 12, 1967. R.E. Schultes 24572 (Alpha-Helix
Amazon Expedition, 1967).

The bitter seeds of Herrania Camargoana are pulverized
and employed as a condiment on game-meat by the Waika
Indians of the Rio Cauaburi in northwestern Brazil.

GUTTIFERAE

Caraipa parvielliptica Cuatrescasas in Rev. Acad. Col. Cienc.

8, No. 29 (1950)64.

CoLomBIA: Comisaria del Vaupes, Rio Kananari, Cerro Isibukuri. “Bush
along rill.”’ April 12, 1951. R.E. Schultes et 1. Cabrera 14738. —
Comisaria del Amazonas, Rio Apaporis, Soratama. ** Height 60
feet.” March 26, 1952. R.E. Schultes et 1. Cabrera 16070. — Rio
Miritiparana. **Small tree. Yukuna: seé-na.”’ May 8, 1952. R.E.
Schultes et 1. Cabrera 16460.

The Yukunas apply the sap of Caraipa parvielliptica to sores
of the mucous membrane of the mouth. In Brazil, the sap of C.
paraensis Huber and C. grandifolia Martius is similarly em-
ployed for herpes, mange and itches (LeCointe: A Amazonia
Brasileira 3(1934)424).

Several species of Caraipa have been reported to contain a
high content of resins which are useful in treating a variety of
skin diseases. A highly toxic vermifugal constituent has been
found in the seeds (Freise, F.W.: Apoth. Zeit. 44(1929) 1481).

233

Symphonia globulifera Linnaeus filius, Suppl. (1781)302.

CoLomBIA: Comisaria del Amazonas, Rio Apaporis, Soratama. ‘*Flowers
red. Height 75 feet."” December 14, 1951. R.E. Schultes et I.
Cabrera 14904.

The bark of this tree, source of a very useful resinous latex, is
said by the Indians of the middle Rio Apaporis to be very
effective, when burned to ashes and applied to recalcitrant
ulcers of the abdomen and legs, in rapidly drying the infection.

Vismia ferruginea Humboldt, Bonpland et Kunth, Nov. Gen.
et Sp. 5(1821)141.
BRAZIL: Estado do Amazonas, Manaos, Flores. ‘‘ Flowers greenish white.

Common bush in scrub growth.’ July 5 - August 12, 1967. R.E.
Schultes 24594 (Alpha-Helix Amazon Expedition 1967).

The resinous exudate of this bush is commonly applied to
sores of the skin in the region of Manaos.

FLACOURTIACEAE

Banara guianensis Aub/et, Pl. Guian. Fr. (1775)548.

COLOMBIA: Comisaria del Vaupes, Rio Vaupés, Mitt and vicinity. ** Bushy
tree, common on river bank. Flowers green-yellow.”’ Septem-
ber 27 - October 20, 1966. R.E. Schultes, R.F. Raffauf et D.
Soejarto 24176.

A spot test with Dragendorff reagent gave a doubtful positive
result for alkaloids.

Mayna longifolia Poeppig et Endlicher, Nov. Gen. et Sp. 3
(1845) 64.
CoLoMBIA: Comisaria del Amazonas, Rio Miritiparana, Cano Guacoya.

‘Fruit cauline, greenish white.’ April 25, 1952. R.E. Schultes et
Cabrera 16285.

The seeds of this shrub are crushed and boiled in water to
make a tea to provoke vomiting in cases of serious food poison-
ing, especially from tainted fish. The tea must, however, be
used with caution, since it is reputedly toxic, causing extreme
dizziness, profuse sweating and uncontrollable trembling.

The plant is well known by all Indians of the area. The

234

Makuna call it o0-too-mee-ko; the Mirana, do-ro-he; the Tani-
muka, ya-poo-moo-ho,; the Yukuna, ka-sd-ra (“beetle tree’’).

Mayna toxica R.E. Schultes in Rhodora 65(1963)16, t. 10.

CoLoMBIA: Comisari del Amazonas, Rio Caqueta, La Pedrera and vicinity,
Quebrada Tonina. On high land. **Small tree, 20 feet tall. Flow-
ers white.” October 5, 1952. R.E. Schultes et 1. Cabrera 17731.

The Mirana Indians of the La Pedrera region assert that
formerly the bark of seeds of this tree were given to dogs as a
poison. The same uses have hitherto been reported (Schultes,
R.E. in Rhodora loc. cit.) from the Vaupes and for other Indian
tribes. It was then indicated that: ‘“*The fact that at least two
species — Mayna muricida and M. toxica — are similarly
employed for their toxic properties by Indians in far-separated
parts of the Colombian Amazon suggests that an investigation
into the chemical constituents of this genus might be of inter-
est.”

Ryania angustifolia(Turcz.) Monachino in Lloydia 12 (1949) 21.

COLOMBIA: Comisaria del Vaupeés, Rio Vaupes, Mitu and vicinity. “Small
tree in secondary growth. Flowers white. Leaves and twigs:
alkaloid doubtful. Bark: alkaloid negative.’’ September 27 -
October 20, 1966. R.E. Schultes, R.F. Raffauf et D. Soejarto
24175,

The root, although not used, is considered poisonous by the
Kubeo Indians.

One species, Ryania dentata Miq., has been reported as an
ingredient of an arrow poison in Colombia (Mezey, K.: Rev.
Acad. Col. Cienc. Exact. Fis. Nat. 7(1947)319.

Sundry species of Ryania are recognized in South America
as poisonous. The toxicity is due presumably to a glycoside
(Merz, K.W.: Arch. Pharm. 268(1930)592; Nakarai, S. et T.
Sano: Arch. Pharm. 272(1943)1).

COMBRETACEAE

Combretum Cacoucia Exe// in Kew Bull. 1931(1931)469.

BRAZIL: Estado do Para, Utinga, Belem. ‘‘Rabo de arara. Extensive liana.

pe)

Acrid water in stem. Flowers red, said to be poisonous.’ Septem-
ber 1947. R.E. Schultes 8668.

There are numerous reports, similar to the folk report from
Belem do Para connected with this collection, that the flowers
of Combretum Cacoucia are toxic. There is apparently no
chemical evidence to sustain this assertion, yet the number of
reports is such that the problem bears serious study.

Caffeine and tannins have been reported from the genus
Combretum (Gibbs R.D.: Chemotaxonomy of Flowering
Plants 3 (1974) 1478).

SOLANACEAE

Saracha procumbens (Cav.) Ruiz et Pavon, Fl. Peruv. 2 (1799)

43.

CoLomBia: Comisaria del Putumayo, Valle de Sibundoy, Sibundoy and
vicinity. Alt. 2225-2300 m. May 29, 1946. R.E. Schultes et M.

Villarreal 7615. — Same locality. August 22, 1963. M.L. Bristol
1328.

According to Bristol, the Kamsa Indian name of this garden
plant, the fruit of which ts edible, is chuftanguemesha. A tea of
the whole plant is drunk as a diuretic and febrifuge (Schultes et
Villarreal 7615).

BIGNONIACEAE

Pseudocalymma alliaceum (Lam.) Sandwith in Rec. Trav. Bot.

Neerl. 34(1937)210.

CoLoMBIA: Comisaria del Amazonas, Rio Amazonas, Leticia. Alt. 100 M.
‘‘Arbusto de 1.5 m., esteril. Olor fuerte aliaceo. Alcaloides:

positivo. Nombre vulgar: sacha-ajo.”’ October 8, 1961. J.M.
Idrobo 4687.

Although this plant is alkaloid-positive with a Dragendorff
test, the whole plant, crushed and made into a tea, is taken
frequently to relieve pulmonary ailments.

236

BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY

VOL. 26, No.7

CAMBRIDGE, MASSACHUSETTS, SEPTEMBER 30, 1978

FOSSIL POLLEN AND THE
ORIGIN OF CORN

ANCIENT POLLEN FROM DEEP CORES IN MEXICO SHOWS
THE ANCESTOR OF CORN TO BE CORN AND NOT ITS
RELATIVE, TEOSINTE.

PAUL C. MANGELSDORF,* ELSO S. BARGHOORN,**

There are currently two main schools of thought on the
question of corn’s origin: one, a 19th century concept, consid-
ers teosinte, corn’s closest relative, to be its ancestor; the
other, a more recent One, maintains that the ancestor of culti-
vated corn was a wild corn, now probably extinct.

The teosinte theorists argue that since the majority of culti-
vated species have extant wild ancestral forms, corn must also
have its wild counterpart and does so in teosinte (/), a species
that is widely distributed in parts of Mexico and Central
America and which has, for more than a century, been recog-
nized as corn’s closest relative (2). This school also relies
strongly on certain parts of the cytogenetic evidence (3), and
this particular evidence is indeed impressive. Corn and teo-
sinte have the same chromosome number, hybridize freely, the
hybrids are usually highly fertile, the pairing of the parental
chromosomes in the hybrids is virtually complete and —
perhaps most important — crossing over between them is
*Fisher Professor of Natural History Emeritus, Harvard University, and Lecturer in

Botany, University of North Carolina.** Fisher Professor of Natural History, Harvard
University.*** Honorary Research Fellow, Botanical Museum, Harvard University.

$25.00 a year, net, postpaid. Orders should be directed to Secretary of Publications at the above
address. Second-Class Postage Paid at Boston, Massachusetts.

essentially the same as it is in pure corn (4). All of these
circumstances combine to make the teosinte theory much more
plausible than it was in the nineteenth century.

The teosinte school concedes that the archaeological evi-
dence presently available lends little support to the teosinte
theory but regards this evidence as being outweighed by the
cytogenetic evidence. It suggests that supporting archaeologi-
cal evidence is lacking because the early stages in teosinte’s
domestication occurred in open campsites where cultural re-
mains have not been preserved (5, 7).

On the question of the fossil pollen discovered in Mexico, the
teosinte school is distinctly ambivalent. On the one hand it says
that this evidence is not to be taken seriously or the data are
confusing and ambiguous and they do not solve the problem of
the origin of corn (5). On the other hand the school asserts that
because of the relevance of the fossil pollen to the validity of
the teosinte hypothesis it requires rigorous examination (6).
That examination results in the rather conflicting conclusions
that the fossil pollen is not large enough to be reliably distin-
guished from teosinte, but is too large to be the pollen of a
primitive wild corn (/9, /, 20). Finally the school explains the
fossil pollen as contamination occurring during the core-
sampling operation (/).

The corn theorists, including the present authors, agree that
cultivated corn undoubtedly had an ancestral form and hold
that this was a wild corn, probably now rendered extinct ini-
tially because of repeated hybridization with cultivated corn
once the practice of agriculture began, and later by the depre-
dations of Old-World grazing animals, horses, cows, burros,
sheep and goats introduced by the Spaniards and other col-
onists. This school sees corn differing from teosinte in numer-
ous genetic, morphological, taxonomic and evolutionary
characteristics (8, 9), and regards the genetic evidence, consid-
ered as a whole, as showing teosinte differing from corn not by
afew genes (/), but by genes or blocks of genes on virtually all
of its chromosomes (8). This school regards the archaeological
evidence as critical and the paleobotanical evidence involving
the fossil pollen as virtually conclusive (/0, /3).

It is the authenticity of the fossil pollen that we wish espe-

238

cially to consider here. Other kinds of evidence bearing on the
problem are treated in detail elsewhere (8, 9). Suffice it to say
here that contrary to a recently published drawing showing an
Indian cultivating teosinte (//), there is presently no evidence
of any kind, archaeological, ethnological, linguistic, ideog-
raphic, pictorial or historical, to show that teosinte was ever
cultivated as a crop by the American Indians.

DISCOVERY OF THE FOSSIL POLLEN

The discovery of the fossil pollen in question was the result
of meticulous studies of palynological samples from cores
taken at the Bellas Artes site in Mexico City in preparation for
the construction of Mexico's first skyscraper, a 43-story build-
ing. (Fig. 1.) These cores were obtained from Dr. Leonardo

T 43rd. storey
7
a‘

Fig. 1. Fossil pollen, identified as corn pol-

len, was discovered in core samples taken at

depths of 69.3 - 70.5 meters in preparation for

Mexico’s first skyscraper. An arrow indi-
cates the level of volcanic clay in which the
pollen occurred. After Zeevaert (28). De-
pending on the criteria used, this level is var- iver

iously dated at 25 to 80 thousand years ago. It
LL

contains no cultural remains. MUYY
VORCOMC CRq
yoicamec clay

se Ya

Fossil Pollen

a

Zeevaert, the engineer in charge of the core sampling, by
Professor Paul Sears of Yale University for palynological
studies to chart climatic changes as they might be revealed by
changes in frequencies of pollen of various species especially
of pines indicating a drier, cooler period: oak and alder a
warmer, moister one and fir a cooler, moister one (/2). In
analyzing the pollen found in the cores, Sears’ associate, Mrs.
Kathryn Clisby, observed in the lower levels of the profile, 69.3
- 70.5 meters, a number of indubitable grass pollen grains that
seemed too large to be identified as those of ordinary grasses.
Thinking that these might be the pollen grains of teosinte, Sears
and Clisby obtained from Mangelsdorf pollen of several var-
leties of teosinte. When it became apparent that some of the
fossil pollen grains were larger than those of teosinte, Sears,
Clisby and Mangelsdorf agreed that these might be pollen
grains of corn and decided to send the cores to Elso S. Barg-
hoorn, a paleobotanist at Harvard, for further study. Barg-
hoorn and his then graduate student Margaret Wolfe made an
intensive study of the fossil grains, macerating out additional
ones from the core centers to eliminate possible surface con-
tamination and comparing them in size and also in the ratio of
the pore diameter to the long axis with pollen of fourteen
varieties of modern corn, three of teosinte and eight of Tripsa-
cum, amore distant relative of corn which, like teosinte, oc-
curs widely in Mexico.

The results of these comparisons show that, although there is
an overlapping in size frequencies between the pollen grains of
corn and those of teosinte, some of the fossil pollen grains are
much too large to be classified as teosinte. We (/3) concluded
that the large fossil pollen grains were almost certainly those of
a wild maize once growing in the Valley of Mexico, well before
the beginnings of agriculture in Middle America, and this es-
sentially established two important facts:

I. Corn is an American plant and not one of Asiatic
origin.

. The ancestral form of cultivated corn was corn
and not teosinte.

i)

The conclusion with respect to corn’s American origin

240

seems to have been generally accepted. At least no recent
articles arguing for an Asiatic orign of corn have come to our
attention.

The second conclusion, that the ancestor of cultivated corn
is corn, was also generally accepted, at least for a period,
especially when it proved to be quite consistent with ar-
chaeological remains of corn, most notably the oldest of these,
uncovered by Richard MacNeish in the Tehuacan Valley of
Mexico (/4). Many students of corn thought that the long and
sometimes acrimoniously debated problem of the origin of corn
had finally been solved.

In the late sixties the old, now long-dormant theory that the
ancestor of corn is teosinte was rather suddenly revived. Most
prominent in its revival was George Beadle, a Nobel Laureate
in Physiology and Medicine and a retired University Chancel-
lor, who, in an interesting personal account, tells his reasons
for reviving the theory (3).

Beadle was soon joined by others, including Walton Galinat,
who believed that the evidence from morphological charac-
teristics which he was then studying outweighed the evidence
from fossil pollen or the archaeological remains in which he had
participated in describing and publishing (/4, 23).

The new devotees of the revived teosinte theory were gener-
ally not initially deterred by the archaeological evidence which
by this time was considerable — or the evidence from the fossil
pollen, both of which were neither consistent nor compatible
with the concept of corn as a domesticated teosinte. Realizing.
however, that the evidence of the fossil pollen was widely
accepted by botanists and archaeologists and could not be
ignored, they tended to dismiss it by relying on Kurtz er al.
(/5). These authors measured pollen from corn plants grown
under a variety of environmental conditions, a treatment which
resulted in considerable variation in the axis/pore ratio, a
measurement which Barghoorn e¢ a/. had earlier employed as
One means of distinguishing the pollen of corn from that of
teosinte. Kurtz er al. concluded that axis/pore ratio alone is not
adequate for making this distinction. The teosinte theory advo-
cates have, with remarkable unanimity, cited or even quoted
Kurtz eal. as raising serious doubts about the identification of

241

the fossil pollen (/, 6, /6) but with equal unanimity have over-
looked or ignored the statement set forth rather conspicuously
in these authors’ summary that their data **do not refute the
findings of Barghoorn et a/.’* and that five of the fossil pollen
grains studied by Barghoorn ef al. “‘are sufficiently large in
both axis length and pore diameter as well as axis/pore ratio to
be classified as maize with a high degree of reliability.”

IDENTIFICATION OF THE FOSSIL POLLEN

The problem of identifying the fossil pollen is one of compar-
ing it with the pollen of corn and its two American relatives
teosinte and Tripsacum. There are no other native grasses with
which the fossil pollen might be identified. Distinguishing the
fossil pollen from that of Tripsacum is not difficult since there is
litte overlapping in size, only the largest grains of Tripsacum
being within the range of the smallest grains of corn. Also the
pattern of spinules on the exines of 7ripsacum grains, as re-
vealed by the scanning electron microscope, is quite different
from that in corn. In Tripsacum the spinules occur in clusters:
in corn they are regularly distributed as they are also in teosinte
(17, 18).

Since the pollen of corn can not be distinguished from teo-
sinte pollen by their spinule patterns which are quite similar,
the only criterion for making a distinction is one of size. It has
been asserted that size is not a taxonomic character. This is not
strictly true. The principal difference between popcorn and
flint corn, for example, is in the size of their kernals. Size of
structures is often included as a part of taxonomic descriptions.

In certain instances there is no difficulty in distinguishing
corn and teosinte pollen by size alone. For example, ina recent
publication (/8), the photographs of pollen of Guerrero teosinte
and Confite Morocho corn show the corn pollen to be only
slightly larger than the teosinte pollen. But when the corn
pollen is enlarged to the same magnification, x 1692, as the
teosinte pollen it proves to be half again as long as the teosinte
pollen with about twice the volume.

Although individual grains of corn pollen cannot always, as

242

was possible in this case, be distinguished from individual
grains of teosinte, populations of pollen grains can usually be
distinguished. A comparison of the frequency distributions of
200 archaeological grains of pollen from the Bat Cave site in
New Mexico with 200 grains of pollen from a teosinte growing
in the Valley of Mexico shows some overlapping (/9). In the
region of overlap the grains of corn and teosinte cannot be
distinguished. But 59 percent of the Bat Cave pollen grains are
larger than the largest teosinte grains. Distinguishing these
from the teosinte grains is no problem.

The population of the fossil pollen grains from the Bellas
Artes site is clearly different from any population of teosinte
grains with which it has been compared. There are, as Kurtz e7
al. have stated, at least five pollen grains, 36 percent of the
total, too large to be identified as teosinte pollen.

The teosinte theorists now argue that the fossil pollen grains,
earlier considered to be too small to be reliably distinguished
from teosinte grains, are too large to be those of a primitive
corn (/, 5). This argument is based on a correlation showing a
relationship between length of ear and pollen size (20). The
length of the ear determines to a considerable extent the length
of the styles, commonly called *‘silks’’, that the pollen tubes
must travel to reach the ovules and effect fertilization.

Since the ears of a primitive wild corn are assumed to have
been small, the earliest intact cobs from San Marcos Cave in
the Tehuacan Valley vary in length from 19 - 25 mm (/4), it Is
concluded that the pollen of such a corn must have been corre-
spondingly small. This does not necessarily follow. Actually
the correlation mentioned above, although statistically signifi-
cant, is strongly influenced by two races, Jala and Huesillo,
that have unusually long ears and unusually large pollen. The
correlation among the remaining eight races included in the
study is by no means so strong and there are notable excep-
tions. The highly evolved Mexican race Vandeno, for example,
has pollen grains of about the same size, 83.9 microns, as those
of the primitive Mexican popcorn race Nal-Tel, 81.2 microns,
although its ears are more than twice as long as Nal-Tel, 17.2
and 7.9 cm, respectively.

243

SIGNIFICANCE OF THE LARGE POLLEN

It is the large pollen of the primitive popcorn that seems to us
to be especially significant and to require explanation. The
length of the mature ear is only one factor in determining the
length of the styles that the pollen tubes must travel to reach the
ovules. Equally important is the extent to which the husks
protrude beyond the tips of the ears which they enclose. The
senior author has frequently been impressed by the fact that
among the early archaeological remains the husks are consid-
erably longer, on the average, than the longest cobs of the
same level. This first came to attention in the archaeological
remains turned up in the first Bat Cave expedition. The only
husk found in the lower levels of the cave is quite long, 24.5cm,
more than twice the length of the longest intact cob, 10.3 cm.
The authors (2/) concluded that the husk must have been an
involucre of leaf sheaths subtending and surrounding the base
of an ear but not tightly enclosing it.

The real nature of this long husk became apparent with a
study of the specimens from the second Bat Cave expedition
(22). These led to the conclusion that the long husk found in the
first expedition may have enclosed, not a single ear, but a pair
of ears, upper and lower, each with its own shorter husk system
(Fig. 2). If the silks of the lower ear became exposed to pollina-
tion only when they reached the terminus of the outer husk
system they would have been about 23 cm long. This is longer
than any of the ears in the correlation study reported above
except the giant ears of the races Jala and Huesillo, but is about
the length of the longest silks of the race Vandeno. when
allowance is made for the husks extending several inches be-
yond the tip of the ear as they do in most varieties. Thus the fact
that the highly-evolved race Vandeno has the same pollen size
as the primitive popcorn race, Nal-Tel becomes explicable.

The two-eared husk system may also explain the two-
ranked, four-rowed cob found in the lower level of San Marcos
cave. The teosinte advocates regard this as evidence of the
evolution of the early Tehuacan corn from teosinte. A more
simple and obvious explanation is that this cob represents a
lower secondary ear in a two-eared husk system. Lower sec-

244

Fig. 2. Archaeological evidence shows that primitive
corn may sometimes have borne two ears, an upper and a
lower, in the same husk system. To reach the terminal

opening of the outer husks the styles — “silks’” — attached
\ to the basal ovules of the lower ear would have to be quite
\ long and the pollen grains effecting fertilization quite large.
| i A This wild corn may have been *‘preadapted”* to evolve

i \ /; under domestication in the direction of producing long
i Yoo 4

single ears. Teosinte does not have this preadaptation.
Solid lines represent actual parts: broken lines artist's re-
construction. Drawn by Julian Camara-Hernandez (22).
]

all

1/2 actual size.

ondary ears, even in modern corn varieties, are often two-
ranked and four-rowed.

Husk systems similar to the one described from Bat Cave
and illustrated in Fig. 2, have also been found in the remains
from San Marcos Cave in Tehuacan and the Huarmey Site in
Peru (23, 24).

Primitive corn as we now conceive of it with its large pollen
may thus be recognized as a classic example of what some
students of the dynamics of domestication, notably Vavilov

245

and Hawkes (25), regard as *‘preadaptation.’’* These authors
stress the fact that primitive ancestral forms of successul culti-
vated plants already possessed tendencies which induced man
to cultivate them. Corn had its share of these especially in its
easily harvested and threshed grain, and its conspicuous re-
sponse to man’s ministrations: freedom from competition with
other vegetation and increased soil fertility. Early cultivators,
although perhaps noting corn’s large pollen grains, just visible
to the naked eye, could scarcely have been aware of their
significance. But it was its large pollen grains that gave corn the
ability to evolve in the direction of producing larger and larger
ears. If ever there was a wild species preadapted to domestica-
tion, corn is perhaps the prime example of this condition.
Teosinte does not have this preadaptation: it could not have
evolved in this direction without a series of mutations involving
pollen size.

POSSIBILITY OF CONTAMINATION

One recent explanation of the fossil pollen is that it repre-
sents contamination (/), either from modern pollen at the upper
levels being carried to a lower one during the sampling or from
pollen in the air when the sampling was being done. To explain
the fossil pollen as contamination is, of course, to assume that
it is indeed corn pollen.

That pollen in the | - 3 meter level could be carried down to
the 69.2 - 70.5 meter level leaving no contamination at interven-
ing levels seems quite improbable, if not impossible, in view of
the precision of the equipment and the techniques employed in

“Further reading has shown us that the idea of preadaptation as presented by these
authors, is new only as it applies to evolution under domestication. Simpson in his
book, Tempo and Mode in Evolution (Columbia University Press, 1944) discusses at
length the concept as it applies to evolution in nature. Students of corn’s evolution,
including the senior author of the present article, have perhaps been remiss in not
recognizing the phenomenon of preadaptation as an important factor in corn’s amaz-
ingly rapid evolution under domestication. Included among corn’s preadaptive charac-
teristics is its ability to hybridize and exchange genes with its relatives, teosinte and
Tripsacum, Teosinte lacks this characteristic, since, if it is cultivated corn’s only
ancestor, it had no wild corn, Zea Mays, with which to hybridize and its does not, even
under ideal experimental conditions, hybridize successfully with Tripsacum.

246

the core sampling. The possibility of contamination by atmos-
pheric pollen appears to be equally remote.

The Bellas Artes site is centrally located in greater Mexico
City, one of the largest metropolitan areas in this hemisphere.
It is many miles removed in all directions from the nearest
corn-growing areas. We do not know exactly how far corn
pollen can be carried by the wind but compared to the pollen of
some other species, pines for example, it is relatively heavy.
Agronomists and seed producers maintaining the purity of
breeding lots of corn allow about one thousand feet of isolation
if the contaminating pollen comes from the windward side, less
if physical barriers to air flow are present (26). During the time
of year that corn pollen would be shedding in Mexico City, if
corn were there, the air is relatively still, indeed so much so
that industrial pollution not carried away by the wind has
become a major problem.

An experiment performed for another purpose by a Harvard
graduate student, Ramana Tantravahi, may have a bearing on
the question of how far corn pollen can travel with the wind. In
order to effect the hybridization of teosinte with 7ripsacum on
a large scale, Tantravahi grew emasculated plants of teosinte
adjacent to pollen-shedding plants of Tripsacum in a small
garden surrounded by University buildings in Cambridge. To
test the effectiveness of the emasculation of teosinte and to
detect contamination by corn pollen from any source he also
grew in the garden arow of emasculated corn plants. When the
ears of these were examined at the end of the season, not a
single kernal was found (27), although there were extensive
plantings of corn in the market-gardening area near the
Waltham Field Station about six miles due west of Harvard
University. The winds during the corn-pollen-shedding season
are prevailingly from the west. These observations suggest that
corn pollen cannot ordinarily be carried as far as six miles.

The possibility that there was corn pollen in the air at the
Bellas Artes site when the soil sampling occurred is quite
remote. Almost equally remote is the possibility that such
pollen, if actually there, could have contaminated the core
samples.

The core sampling at the Bellas Artes site that revealed the

247

fossil pollen is not an ordinary drilling operation involving
repeated pouring of water or drilling mud into the hole, thus
providing abundant opportunities for contamination. Core
sampling in preparation for the construction of a skyscraper is
an operation of considerable precision — one that is especially
designed to yield undisturbed samples. Essentially it involves a
series of tubes driven through the soil at successive levels. As
the tubes are removed, they are immediately sealed at both
ends with a special wax and are sent to the laboratory for
various analyses. Engineers familiar with the problems of soil
mechanics who have examined the data in Dr. Zeevaert’s
article in Geotechnique (28) describe his operation as *‘ex-
traordinarily careful and meticulous.”

Dr. Zeevaert himself seems quite certain that the cores in
which the fossil pollen was found are undisturbed samples. Ina
letter of October 17, 1973, to Barghoorn, he wrote:

Indeed the sampling of the material was performed with a special
sampler to obtain undisturbed samples of the soil, useful to
determine the natural compressibility and sheer strength prop-
erties of the materials. Therefore, the samples taken were not
disturbed or contaminated, they were ‘undisturbed samples’
used in soil mechanics to determine the ‘in situ’ mechanical
properties of the materials. Therefore, you can be sure that the
investigations made on these samples concerning the fossil
maize pollen are reliable.

We are inclined to accept Dr. Zeevaert’s statement as factual.
We are pleased to note, in passing, that Dr. Zeevaert has
recently been elected a foreign member of the United States
National Academy of Engineering.

Finally the fossil pollen itself demonstrates that it is not the
product of modern contamination. It does so in this way: it Is
one of the standard palynological techniques to prepare pollen
for electron microscope studies by a treatment known as
acetolysis (glacial acetic acid and concentrated sulfuric acid,
9:1). This treatment is described in detail by Banerjee (29).
Suffice it to say here that corn pollen when fresh resembles in
shape an inflated basketball: when dry the shape of a deflated
ball. Pollen from extant corn plants is restored to its original
inflated shape when subjected to the acetolysis treatment
(Plate I8A). But the pollen from the Bellas Artes does not

248

respond as does modern pollen to this treatment (Plate 18D).
This is true also of certain other ancient pollen. Pollen from
Coxcatlan Cave in the Tehuacan Valley dated at ca. 1900 years
failed to expand completely with the acetolysis treatment
(Plate 18C). On the other hand pollen from the same cave at ca.
1600 years expanded almost like modern pollen (Plate 18B).
Apparently as the chemical constituents of the pollen grains
change with age certain of them lose their ability to react with
the chemicals introduced by the acetolysis treatment (Plate
I8C). What these constituents might be is a question beyond
the scope of this discussion. The important point here is to
recognize the fact that ancient pollen grains differ from modern
ones in their capacity to respond to certain chemical treat-
ments. The loss of this capacity seems to begin at about 2000
years; we do not know at what age it is completely lost. Pollen
from the Huarmey site in Peru, dated at 3600-4000 years (20) is
only slightly more collapsed than the pollen shown in Plate 18C
dated at 1900 years (29).

The failure of the Bellas Artes pollen to respond to the
acetolysis treatment combined with the fact that opportunities
for contamination, discussed above, are minimal, if not ac-
tually nonexistent, has persuaded us that this pollen is indeed
ancient and not the product of modern contamination. Having
previously satisfied ourselves that the pollen has been cor-
rectly identified as corn pollen, we can now only conclude that
the fossil pollen is authentic and if so, it is more than highly
probable that the ancestor of cultivated corn was corn and not
teosinte.

IMPLICATIONS FOR CORN IMPROVEMENT

Eliminating teosinte as the ancestor of corn does not mean,
however, that it has had no role in the evolution of cultivated
corn. On the contrary, archaeological remains are consistent in
showing that although the earliest corn may have been pure
corn, later corn is the product of hybridization with teosinte.
And the hybridization still continues (30).

To the puzzled onlooker the distinction between recognizing

249

teosinte as an ancestor of modern corn while vigorously deny-
ing it to be sie ancestor may seem to be no more than an
exercise in semantics. From the standpoint of theoretical gen-
etics and practical plant breeding, however, it is much more
than that. If teosinte is fe ancestor of corn and the only one,
then modern corn contains only one stream of germplasm. But
if the ultimate ancestor of corn was corn, as the fossil pollen
and the archaeological remains show, and modern corn Is the
product of repeated introgression from teosinte, then two dis-
tinct streams are involved and modern corn, although a diploid,
has some of the attributes of an allopolyploid (8). In this respect
it is comparable to the allopolyploid cereals like bread wheat,
one of the world’s most productive food plants, the product of
hybridizing three species, two of which have never been rec-
ognized as worthy of cultivation and a third, einkorn, which
being generally quite unproductive, is but little grown.

The interactions of these two streams of germplasm, corn’s
and teosinte’s, has produced profound effects, including gen-
etic recombination, heterosis — hybrid vigor — and mutagene-
sis (3/). These, combined with corn’s preadaptation to changes
in ear length, by virtue of its large pollen, have resulted in the
explosive evolution (23) illustrated in Plate 19. It is doubtful if
such rapid evolution could have occurred were teosinte the
only ancestor of corn. Then the hybridization of cultivated
corm, originating from teosinte, with its ancestor, teosinte
would have produced a minimum of genetic recombination and
heterosis and probably no mutagenesis, and any increase in ear
length would have been inhibited by teosinte’s lack of preadap-
tation because of its small pollen.

To the practical plant breeder the difference in the two
theories may determine the most promising methods of
employing genes from teosinte. Shall he hybridize corn directly
with teosinte or shall he hybridize strains of one race of corn
containing teosinte germplasm with strains of another race
containing a somewhat different combination of teosinte
genes? In both strains the teosinte genes or blocks of genes will
have been absorbed into the corn genotype by the natural
selection of modifying genes that suppress the undesirable
characters of teosinte and allow the desirable ones to be ex-

250

pressed. The genotype of modern corn may well be a constella-
tion on a grand scale of DNA recombinants.

So far little progress in corn breeding has been made by
hybridizing corn directly with teosinte but phenomonal results
have been obtained by combining strains that are the product of
past introgression from teosinte as shown by anatomical and
genetic studies (32).

The average yields of corn in the United States have in-
creased from 26 bushels per acre in 1929 to 95 bushels in 1972.
This progress has involved the bringing together by empirical
methods of strains of germplasm from diverse sources, much of
it originally from teosinte. There are many races of corn not yet
employed in hybrid corn breeding (33). Recognizing the poten-
tial value of these for breeding opens new possibilities for
improvement that may render corn, this nation’s basic food
plant, still more important on the world scene as one of not
more than about twelve species of cultivated plants, each one a
unique biological system, that quite literally stand between
mankind and starvation.

SUMMARY

Fossil pollen discovered in core samples from a depth of 69.3
- 70.5 meters in preparation for the construction of Mexico's
first skyscraper has been identified as the pollen of a primitive
wild corn. Its authenticity seems now to be well established
since the possibility that the pollen represents modern con-
tamination is shown to be remote. The authors conclude that
the ancestor of cultivated corn was a wild corn and not its
closest relative, teosinte, and that this may have important
implications for corn’s genetic improvement and its role in
meeting the world food problem.

POSTSCRIPT
Two important events occurred while this paper was in pro-

of. Anarticle by Iltis et a/. (appearing in Science, Jan. 12, 1979)
251

announced the discovery in Mexico of a diploid perennial
teosinte. As senior author of the present article. I recognized
this as a significant discovery and I wrote Iltis at once con-
gratulating him. I failed, however, to recognize the full signifi-
cance of the discovery. It remained for H. Garrison Wilkes, the
author of the book, Teosinte: the Closest Relative of Maize (2).
presently in India, to point out in a letter to me, dated Jan. 17,
that this discovery may be the key piece in the puzzle, a
so-called **missing link’’ in corn’s genealogy. Wilkes assumes,
correctly I think, that hybridization between the diploid peren-
nial teosinte and a wild annual corn could have produced all of
the known annual races of teosinte. This assumption is to a
large extent testable.

I am urging Wilkes to publish this concept and its implica-
tions as soon as possible, and I am hoping that this postscript
will serve to establish his priority for an imaginative and impor-
tant new idea. The relevance of our present article to this new
concept Is obvious.

REFERENCES AND NOTES

1. G.W. Beadle, in World Anthropology, General Editor Sol Tax Origins
of Agriculture, Charles A. Reed, Ed. (Mouton Publishers, The Hague-
Paris, 1977), p. 615: /bid., in Report of Thirty-Second Annual Corn and
Sorghum Research Conference, H.D. Loden and D. Wilkinson, Eds.
(American Seed Trade Association, Washington, D.C., 1977), p. 1.

2. H.G. Wilkes, Teosinte: the Closest Relative of Maize. (Bussey Inst.
Harvard Univ., Cambridge. Mass., 1967).

3. G.W. Beadle, Field Mus. Nat. Hist. Bull., 43, 2 (1972).

4. R.A. Emerson and G.W. Beadle, Zeitsch. Ind. Abstamm. Vererb., 62,

305 (1932).

5. W.C. Galinat, in Corn and Corn Improvement, G.F. Sprague, Ed.

(Amer. Soc. of Agronomy, Madison, Wisconsin, 1977), p. |.

Ihid., Ann. Rev. Genetics, 5, 447 (1971).

Ibid., in Genetics and Breeding of Maize, D.B. Walden, Ed. (Wiley &

Sons. New York. 1978). p. 93.

8. P.C. Mangelsdorf. Corn Its Origin, Evolution, and Improvement. (Har-
vard Univ. Press, Cambridge, Mass., 1974).

9. L.F. Randolph, Econ. Bor. 30, 321 (1975).

10.) P.C. Mangelsdorf, Field Mus. of Nat. Hist. Bull., 44, no. 3, 16, (1973).

ll. H.G. Wilkes, Science. 200, 41 (1978).

12. K.H.Clisby and P.B. Sears, Bull. Geo. Soc. Amer., 66, 511 (1955): P.B.
Sears and K.H. Clisby. /bid., 521 (1955).

aS

252

E.S. Barghoorn, M.K. Wolfe. K.H. Clisby, Bot. Mus. Leafl. Harvard
Univ., 16, 229 (1954); P.C. Mangelsdorf, Amer. Antiquity, 19, 409
(1954).

P.C. Mangelsdorf, R.S. MacNeish, W.C. Galinat, Science, 143, 538
(1964).

E.B. Kurtz, J.L. Liverman, H. Tucker, Bull. Torr. Bot. Club, 87, 85
(1960).

J.M.J. De Wet and J.R. Harlan, Euphvtica. 21,27(1972): T.A. Kato Y.,
Cytological Studies of Maize and Teosinte in Relation to their Origin
and Evolution. (Mass. Agric. Exper. Station, Univ. of Mass. Amherst,
1975): J. Schoenwetter, Amer. Antiquity. 39, 292 (1974).

U.C. Banerjee and E.S. Barghoorn, 30th Ann. Proc. Electron Micro-
scopy Soc. Amer., C.J. Arceneaux, Ed. (1972).

C.A. Grant, Grana. 12, 177 (1972).

W.C. Galinat, Maize Genet. Coop. Newslet., 47, 105 (1973).

W.C. Galinat, Maize Genet. Coop. Newslet., 35, 39 (1961).

P.C. Mangelsdorfand C.E. Smith, Jr., Bot. Mus. Leafl. Harvard Univ.,
13, 213 (1949).

P.C. Mangelsdorf, H.W. Dick, J. Camara-Hernandez, Bot. Mus. Leafl.
Harvard Univ., 22. | (1967).

P.C. Mangelsdorf, R.S. MacNeish, W.C. Galinat, in The Prehistory of
the Tehuacan Valley: Environment and Subsistence, D.S. Byers, Ed.
(Univ. of Texas Press, Austin & London, 1967), 1, p. 178.

A. Grobman, D. Bonavia, D.H. Kelley, P.C. Mangelsdorf, J. Cam-
ara-Hernandez, Bot. Mus. Leafl. Harvard Univ., 25, 221 (1977).
J.G. Hawkes, in The Domestication and Exploitation of Plants and
Animals, P.J. Ucko and G.W. Dimbleby, Eds. (Gerald Duckworth &
Co., London, 1969), p. 17; N.I. Vavilov, Bull. Appl. Bot. Pl. - Breed.,
16, 1 (1926).

W.F. Craig, inCorn and Corn Improvement, G.F. Sprague, Ed. (Amer.
Soc. of Agronomy, Madison, Wis., 1977), p. 671.

R. Tantravahi, unpublished.

L. Zeevaert, Geotechnique. 7, 115 (1957).

U.C. Banerjee, Morphology and Fine Structure of the Pollen Grains of
Maize and its Relatives. (Ph.D. Thesis, Harvard Univ., Cambridge.
Mass., 1973).

H.G. Wilkes, Econ. Bot., 31, 254 (1977).

P.C. Mangelsdorf. Cold Spring Harbor Symp. Quant. Biol., 23, 409
(1958).

G.S. Johnston, Manifestations of Teosinte and ‘Tripsacum’’ Intro-
gression in Corn Belt Maize. (Bussey Inst. Harvard Univ., Cambridge,
Mass., 1966); P.C. Mangelsdorf, Euphivtica, 10, 157 (1961): S.M. Seh-
gal, Effects of Teosinte and Tripsacum™ Introgression in Maize. (Bus-
sey Inst. Harvard Univ., Cambridge, Mass., 1963): S.M. Sehgal and
W.L. Brown, Econ. Bot., 19, 83 (1965).

M.M. Goodman and R. McK. Bird, Econ. Bot., 31, 204 (1977): E.J.
Wellhausen, Ann. Hybrid Corn Industry - Res. Conf. Proc., 20, 45
(1965).

Our thanks to Major M. Goodman, L.F. Randolph, Robert Bird, Clark
P. Mangelsdorf, Paul C. Mangelsdorf, Jr., and Dennis Porter.

253

PLATE 18

Plate 18. The effect of the acetolysis treatment on restoring pollen to its original
shape. A. Modern pollen grain of the Mexican popcorn race Polomero Toluqueno, fully
expanded after treatment: B. Archaeological pollen grain from the Tehuacan site,
Mexico ca 1600 years old, expanded: C. Pollen grain ca 1900 years old from the same
site, slightly collapsed; D. Fossil pollen grain from the Bellas Artes site, Mexico,
completely collapsed, showing that it is ancient and does not represent contamination
of the core samples.

254

PLATE 19

cM. | 2 3 4 5 6 7 8 9 10

Plate 19. Evolution in ear length in 6500 years in the Tehuacan Valley, Mexico. The
approximate ages of the cobs from left to right are 7000, 5300, 2200, 450 and 450 years
respectively (23). This evolution, rapid as compared to that of other cultivated plants,
is partly the product of wild corn’s preadaptation to domestication by virtue of its large
pollen as revealed by the fossil remains from the Bellas Artes site, Mexico.

255

BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY

CAMBRIDG

E, MASSACHUSETTS, OctoRER 31, 1978 Ve 26, No. 8
EFFECTS OF FIELD PRESERVATION ON THE
FLAVONOID CONTENT OF JESSENIA BATAUA

GILLIAN A. COOPER-DRIVER* AND MICHAEL J. BALICK**

As a preface to a chemotaxonomic investigation on the Jes-
senia-Oenocarpus complex (Palmae), it seemed desirable to
determine how specimens of plants collected in the field in the
wet tropics should be best preserved for future chemical inves-
tigation.

Up to 30-40 years ago, most collected plant material was
dried by placing it in a conventional press with frequent change
of blotters to absorb moisture from the specimens, the entire
process taking about a week (1). However, in the wet tropics
much of the material was damaged by insects or micro-organ-
isms before drying was complete, rendering the specimen of
little value for taxonomic purposes. The artificial drying of
such specimens by the use of field stoves or even electric bulbs
overcame these problems, but collectors could not easily
transport such equipment to remote study areas. Thus, the use
of chemical preservatives was introduced to treat plants col-
lected in the field and to prevent them from deterioration by
insects or by rotting, until they could be dried at a base facility.
Schultes (2) suggested a formalin (40% formaldehyde) and
water mixture of approximately 1:5 to avoid the spoilage of
most plant materials. Moore (3) recommended using a 1%
aqueous solution of hydroxyquinoline sulfate, while Hodge (4)
cited the use of an ethanol mixture, noting also that the use of

*Department of Biological Sciences, Boston University, Boston, Massachusetts.
*“Botanical Museum of Harvard University, Cambridge, Massachusetts.

Botanical Museum Leaflets (USPS 404-990). Published monthly except during July and A ugust by
the Botanical Museum, Harvard University, Cambridge, Massachusetts 02138. Subscription:

$25.00 a year, net, postpaid. Orders should be directed to Secretary of Publications at the above
address. Second-Class Postage Paid at Boston, Massachusetts.

potable spirits worked quite well if nothing else was available.

Although it is certain that such methods are useful for the
adequate preservation of plant specimens for classical taxo-
nomic purposes, it appeared likely to us that specimens pre-
served in such ways might have suffered a change in their
chemistry. For example, formaldehyde is known to react with
many aromatic compounds, while treatment with aqueous al-
cohols (ethanol and methanol) is the method of choice for
extracting a wide variety of secondary compounds, used in
chemotaxonomic studies, from plant tissues. Consequently, the
use of such methods of preservation might make it difficult to
properly survey the many different classes of such compounds
which have been used for systematic purposes (5). Flavonoids,
for example, as well as related phenolics, might be severely
affected. These are probably the most useful class of secondary
plant constituents for chemotaxonomic purposes because of
their widespread distribution, diversity of structure, and rela-
tive ease of identification using simple apparatus (6).

Since the flavonoids have previously been shown to be use-
ful in the taxonomy of the Monocotyledonae, including the
Palmae (7), they would appear to be a useful group of com-
pounds to throw light on specific problems relating to the
taxonomy of the Jessenia-Oenocarpus complex. Since most of
the material studied grows in the Amazon Valley, however,
many miles from any centers of civilization, it was decided to
determine whether the use of plant preservatives in the field
would affect subsequent analysis of the material for flavonoid
compounds. Preservative treatments similar to those that
might be used in the field were, therefore, investigated under
laboratory conditions, and the results are presented here.

The plants of Jessenia Bataua (Mart.) Burret used in this
study were grown from a population of seeds collected from a
single tree in Tukunare, near Mitu, Comisaria del Vaupés,
Colombia, in July of 1976. The seeds were germinated, planted
in soil and grown under controlled conditions in a phytotron for
one year, until the plants were 20 cm. high. Pinnae were re-
moved and subjected to the following treatments in order to
simulate the procedures that have been used previously in the

258

field (2,4): 1. sun dried; 2. dried in an herbarium oven at 10S°F;
3. soaked in water: 4. soaked in 95% ethanol; 5. soaked in white
rum (86 proof, ca. 44% ethanol); 6. soaked in an FAA mixture
(ethanol 90%: glacial acetic acid 5%: formalin 5%): 7. soaked in
formalin (100% formalin, ca. 40% formaldehyde) and water ina
1:1 mixture; 8. soaked in formalin alone.

The material for treatments 3-8 was stored in the dark for
four weeks at room temperature, and the leaves were then
removed from the solutions and air dried. The solutions were
kept in a refrigerator.

Two samples of dried leaves from each of the eight treat-
ments described above were finely macerated, extracted with
boiling 80% methanol, and the solution concentrated and made
up to a final volume of 10 ml. The ultra-violet spectrum (220-
400 nm) of the methanol extract (suitably diluted) was deter-
mined and the amount of phenolic compounds calculated,
using caffeic acid (a cinnamic acid) and rutin (quercetin 3-0-
rhamnoglucoside, a flavonol) as standards. Identical amounts
of the methanol extracts were also examined by 2-dimensional
chromatography, using the solvent systems n-butanol-acetic
acid-water 6:1:2 (BAW) followed by 5% aqueous acetic acid
(HOAc).

The solutions in which the leaves had been soaked were
concentrated to a standard volume and subjected to 2-dimen-
sional chromatography by the procedure described above.

The amount of phenolic compounds remaining in the leaf
material following these various treatments is shown in Table 1.

TABLE |

Treatment phenols* mgm/gm dry wt.
1. Sun dried 12.5
2. Herbarium dried 11.5
3. Water 3.0
4. 95% Ethanol 2.4
5. White rum 2.4
6. FAA 0.9
7. Formalin-water 1:1 1.8
8. Formalin 1.6

“as caffeic acid

259

Two dimensional chromatography of the sun dried Jessenia
Bataua leaf material gave the following pattern of major pheno-
lic compounds. (Figure 1)

BAW 7
oo
° ; (2)
~- Z
eee
HOAc
FIGURE |. 2-dimensional chromatogram of ultra-violet fluorescent spots of flavo-

noids and related compounds in Jessenia Bataua.

From their fluorescence under ultra-violet light and their
position on the chromatogram, it is believed that compounds
1-3 are flavonol glycosides: 4-5 flavone glycosides: 6 an un-
known cinnamic acid and, compounds 7-8 isomeric forms of
caffeic acid and p-coumaric acid derivatives respectively. Fur-
ther investigations on the identity of these compounds is at
present underway.

The 2-dimensional paper chromatograms showed that the
following compounds were present in the various leaf extracts
(Table 2).

The 2-dimensional paper chromatograms of the solutions in
which the leaves had been soaked showed that the following

260

compounds had been extracted from the leaves and were pre-
sent in the solution (Table 3).

TABLE 2

Treatment Phenolic compounds present

| 2 3 4 5 6 7 8
1. Sun dried ++ ++ ++ +4 ¢4+ ++ F+4+ F4+
2. Herbarium dried ++ ++ ¢4+ 44 44+ +4 F¢4+ +4
3. Water + + + + + + +
4. 95% Ethanol tr tr tr tr tr tr tr tr
5. White rum tr tr tr tr tr tr tr tr
6. FAA ~ = - - = _
7. Formalin-water I: 1 = _ = = = 7 =
8. Formalin —_ i. _ = _ =

(++ = very positive: + = positive; tr = trace: — = negative)
TABLE 3

Treatment Phenolic compounds present

l 2 3 4 5 6 7 8
3. Water + + + + + + + +
4. 95% Ethanol te a Se ee ee ee Sa Oe Se Ce oe
5. White rum ++ t+ +4 ++ 4¢4+ ++ ++ FF
6. FAA + + + + + + + +
7. Formalin-water 1:1 - - - - -
8. Formalin - = = _ _ =

As we expected, there is no doubt from the above results
(Tables 1-3) that, if plant material is to be subjected to chemo-
taxonomic investigation for flavonoids or closely related
phenolic compounds, the material must be preserved only by
sun drying or by subsequent artificial drying. Soaking in pres-
ervatives containing 95% ethanol, white rum and FAA results
in changes of cell permeability and consequent leakage of
phenolic material into the solution (Table 3). Preservatives
containing formaldehyde in high concentration (nos. 7 & 8),
which cross link with proteins, carbohydrates and phenolic
compounds to give insoluble precipitates, show no free pheno-
lic compounds either in methanolic leaf extracts or in the
solutions themselves. Water affects the concentration of the

261

phenolics in the leaf less, but it is not a good preservative
because of its lack of fungicidal or bacteriocidal properties.
Therefore, care should be taken when using tropical herbarium
material, since, in many cases, the nature of their treatment
after collection is unknown.

On the basis of these results, ways of treating of Jessenia
Bataua for subsequent chemical analysis can be rated as fol-
lows: (Table 4; highest rating is 1)

TABLE 4
Treatment Rating
Sun dried |
Herbarium dried
Water 2
95% Ethanol 3
White rum :
Formalin-water 1:1 4
Formalin
FAA 5

Based on our preliminary experiments, the following sug-
gestions can be made:

First: All herbarium material should be plainly marked on
the label as to whether it has been chemically treated
in the field and if so for how long and in what way. If
specimens have been treated they are occasionally
so marked, but rarely is it indicated that specimens
were merely sun dried or herbarium dried. This lack
of information reduces the value of these specimens
as potentially useful samples for chemical study.

Second: With the increasing use of chemotaxonomy as one of
the tools for systematics, at least 10 gms. and prefer-
ably 50 gms. of leaf material (or flowering material,

262

or both depending on the plant) should, whenever
possible, be collected, sun or ‘blotter’ dried separate-
ly, and affixed in an air tight bag to the original
specimen for later chemotaxonomic use. This is es-
pecially urgent for tropical material, since the
habitats listed on herbarium sheets often no longer
exist. As flavonoids have been identified in her-
barium specimens over 100 years old, there is little
problem of subsequent shortage! A few hours of
extra field effort in caring for this material might then
help towards solving taxonomic problems which
could arise in the future, especially during expedi-
tions of ‘‘salvage botany’’, collecting from important
localities due to be destroyed by colonization or
programs of *‘development through destruction’.
Obviously, further studies are needed on the use of preser-
vatives covering the entire range of tropical material, including
monocotyledyons and dicotyledons, ferns and lower plants, as
has in part been done by Giannasi for temperate plants (pers.
comm.). With chemotaxonomy less an experimental subject
and coming of age as a valid systematic tool, our field methods
must be readjusted to provide a source of reliable material for
the future investigator. Thus, the use of this material will, it is
hoped, serve in our attempt to understand the complex and
dynamic biological order of the tropics.

ACKNOWLEDGMENTS

For valuable comments and assistance in preparing this pa-
per, we are grateful to Prof. Richard Evans Schultes and Mr.
Michael Donoghue of Harvard University, and to Prof. Tony
Swain of Boston University. Mr. Kim Laver, a student at
Boston University, assisted in the experimental work. Funding
from the following sources for field work in South America is
gratefully acknowledged: Centro de Desarrollo Integrado
“Las Gaviotas”, Colombia, Dr. Paolo Lugari Castrillon, Di-
rector; Sigma-Xi, the Scientific Research Society of North
America; the Atkins Fund of Harvard University.

263

to

LITERATURE CITED

. Lawrence , G.M., 1951, Taxonomy of Vascular Plants, The MacMillan

Co., New York.

Schultes, Richard E., 1947, **The Use of Formaldehyde in Plant Collect-
ing’ Rhodora 49:54-60.

Moore, H.E., Jr., 1950, °°A Substitute for Formaldehyde and Alcohol in
Plant Collecting’? Rhodora 52:123-124.

Hodge, W.H., 1947, °**The Use of Alcohol in Plant Collecting’ Rhodora
49:207-210.

. Bendz, G. and Santesson, J. (Eds.), 1974 Nobel 25, Chemistry in Botant-

cal Classification, Academic Press, London.

Harborne, J.B. in The Flavonoids, 1975, (J.B. Harborne, T.J. Mabry and
H. Mabry Eds.) Chapman and Hall, London p. 1050.

Williams, C.A., Harborne, J.B. and Clifford, H.T., 1973, “‘Negatively

Charged Flavones and Tricin as Chemosystematic Markers in the
Palmae’™> Phytochemistry 12:2414-2430.

264

PLATE 20

Py ae

wegne

t
33
:
a
“

~

y me,
BK a ce gt

Plate 20. Jessenia Bataua (Mart.) Burret inaswampy habitat near Manaus,
Brazil. - Photograph by M.J. Balick.

265

BOTANICAL MUSEUM LEAFLETS VOL. 26, No. 8
OcTOBER 31, 1978

DE PLANTIS TOXICARIS E MUNDO
NOVO TROPICALE COMMENTATIONES XXV

BIODYNAMIC ACANTHACEOUS PLANTS
OF THE NORTHWEST AMAZON

RICHARD EVANS SCHULTES

Continuing a series of articles to call attention to ethnophar-
macologically important plants of the northwest Amazon, I
herewith present notes on uses of several acanthaceous species
which, during my field work in Colombia from 1941 through
1954, I encountered in the local native pharmacopoeias.

Most of the collections were identified by the late Dr. Emery
C. Leonard of the Smithsonian Institution and have been cited
in his The Acanthaceae of Colombia in Contributions of the
United States National Herbarium 31. pt. 3 (1953).

Little is known of the phytochemistry of the Acanthaceae.
Traces of an alkaloid have been found in Justicia gendarussa
L. (Hegnauer, R.: Chemotaxonomie der Pflanzen 3(1964)46).
Saponines have been reported from Aphelandra, Justicia,
Mendoncia and Sanchezia; tannins from Aphelandra and San-
chezia: and cyanogenesis from Aphelandra aurantiaca, Fit-
tonia Verschaffeltii, Justicia carnea and Sanchezia nobilis.
(Gibbs, R.D.: Chemotaxonomy of Flowering Plants 3(1974)
1774-1775).

Specimens of the collections cited are preserved in the
United States National Herbarium; the Gray Herbarium or the
Economic Herbarium of Oakes Ames at Harvard University;
and the Herbario Nacional de Colombia in Bogota.

Aphelandra aurantiaca (Schiedw.) Lindley in Bot Reg. 31
(1845) t.12.

COLOMBIA: Comisaria del Amazonas, Trapecio Amazonico, Rio Loreto-
yacu *‘Herb. Flowers vermillion’’. October 20-30, 1945. R.E. Schultes 6630.

267

The root bark of Aphelandra aurantiaca is considered by the
Tikuna Indians of the Trapecio Amazonico to have strong
properties when administered in a warm decoction to allev-
late progressive deafness due to old age. The decoction is
dropped into the ears.

Aphelandra pilosa Leonard in Contrib. U. S. Nat. Herb. 31
(1951) 209.

CoLomBIA: Comisaria del Vaupes, Rio Guainia. ** Flowers scarlet’. Novem-
ber 17, 1952. R.E. Schultes, R.E.D. Baker et 1. Cabrera 18455.

A cold poultice of the crushed leaves of Aphelandra pilosa is
applied by the Kuripako Indians of the Rio Guainia to relieve
swellings of the throat, due probably to chronic inflamation of
the amygdalae.

Fittonia argyroneura £. Coemans in Fl. des Serres 15 (1865)
103.

EcuaApbor: Napo, Rio Aguarico, Dureno. **Primary forest’’. December 21,
1965. H. Pinkley 15.

According to the collector, this small plant, which the Kofan
Indians call minakoro - sé - hé - pa, is made into a tea which is
used to rinse out the mouth to relieve toothache.

Fittonia Verschaffeltii (e77.) &. Coemans in Fl. des Serres 15
(1865) 186).

CoLomBIA: Comisaria del Putumayo, Rio Sucumbios, Quebrada Conejo,
April 2-5, 1942. R.E. Schultes 3480.

The Kofan Indians, who call this herb mee-nd-ku-ru, state
that it is used medicinally to alleviate a condition probably
allied to chronic nephritis. A tea of the leaves is drunk three or
four times a day when there is difficulty or pain in urination,
usually in elderly members of the tribe. The decoction is also
taken for sore throat and as a febrifuge.

Justicia Blackei Leonard in Contrib. U. S. Nat. Herb. 31 (1951)
SO8.

CoLOMBIA: Comisaria del Amazonas, Trapecio Amazonico, Rio Loreto-

268

yacu. ‘‘Herb. Flowers white and purple.’’ September 1946. R.E. Schultes et
G.A. Black 8317.

The Tikuna Indians of the Trapecio Amazonico, who know
this plant as ké-ré=ma, valued the leaves in infusion to relieve a
condition which seems to be due to chronic sinusitis. The tea 1s
gargled and drunk. The Tikunas appear to be especially prone
to sinus-related affections in the wet season.

Justicia Cabrerae Leonard in Contrib. U. S. Nat. Herb. 31
(1951) 566.

CoLomsBia: Comisaria del Amazonas, Rio Pacoa and Rio Kananart. ** Flow-
ers white.’ June 20, 1951. R.E. Schultes et 1. Cabrera 12701.

The roots of Justicia Cabrerae are esteemed by the Makuna
Indians as the basis of a remedy for skin affections, probably of
a fungal origin, of the crotch. The powdered root is mixed with
lard from boars and rubbed repeatedly on the affected area.
The Makuna refer to the plant as mee-kee-na-ta-ree.

Justicia chlorastachya Leonard in Contrib. U.S. Nat. Herb. 31
(1951) 495.

Cotomsia: Comisaria del Amazonas, Rio Apaporis, Soratama. June 20,
1951. R.E. Schultes et I. Cabrera 12703.

The pulverized roots of this plant are employed in the same
manner as those of Justicia Cabrerae. The Makuna name is the
same.

Justicia comata (L.) Lamarck Encycl. 1(1785)632

CoLomBiA: Comisaria del Amazonas, Rio Hamacayacu, between Amazon
and Putumayo watersheds. **Herb. Flowers white, violet-spotted.’” Septem-
ber, 1946. R.E. Schultes 8248.

The Tikuna Indians gather the plants of Justicia comata, dry
the leaves and pulverize them. Not only are the powders em-
ployed as perfumes, but they are believed to be efficaceous
insect repellants when dusted on the body.

Justicia ideogenes Leonard in Contrib. U. S. Nat. Herb. 31
(1251) G22.

269

CoLomMBIA: Comisaria del Putumayo, Rio Sucumbios, Quebrada Conejo.
April 2-5, 1942. R.E. Schultes 3538.

Justicia ideogenes is one of the Kofan Indian remedies for a
condition leading to a palsy-like trembling, usually associated
with age. The whole plant is boiled in water, and the drink is
poured warm with friction over the lower arms and the feet.
The Kofan name of J. ideogenes is mee-ré-kd-sé-pa and chu-
ru-ko-pu.

Justicia pectoralis Jacquin, Enum. Pl. Carib. (1760) 11.

BRAZIL: Territorio do Roraima, Posto Mucajai, Rio Mucajat. “Secondary
forest by airstrip. Herb. Petals blue, stamens white.’’ March 21, 1971. G.T.
Prance, W.C. Steward, J.F. Ramos et O.P. Monteiro 111 74.

According to the collectors, this plant is ‘used by Indians for
hallucinogenic & snuff’. It is called by the Uaicd-Mucajai
paxararok,

Justicia pectoralis Jacquin var. stenophylla Leonard in Contrib.
U.S. Nat. Herb. 31(1951)615.
BRAZIL: Estado do Amazonas, Rio Cauaburi, Maturaca. Waika=mashihiri.

July 5- August 12, 1967. R.E. Schultes 24573 (Alpha-Helix Amazon Expendi-
tion, 1967)
Territorio do Roraima, Rio Tototobi, Waika village of Wayhana-oo-thle,

August 6, 1967. R.E. Schultes 24627 (Alpha-Helix Amazon Expedition,
1967).

Justicia pectoralis var. stenophylla is extensively cultivated
in Waika villages. The leaves are dried, pulverized and added
to the powdered “resin” of Virola theiodora (Spr. ex Benth.)
Warb. in the preparation of the hallucinogenic snuff known in
the region as epena ornyakwana (Schultes, R.E. and B. Holm-
stedt: Rhodora 70( 1968) 113-160).

It is possible, even probable, that this plant — as well as
other species of Justica — may be used alone, without admix-
ture, by various groups of the Waika in Brazil and Venezuela as
the source of an hallucinogenic snuff (Schultes, R.E. in Efron,
D. [Ed.] Ethnopharmacologic Search for Psychoactive Drugs
(1967) 303; Chagnon, N., P. Le Quesne and J. Cook: Acta
Cient. Venez. 21(1970)186-193).

270

It is clear that much more field work must be carried out
before the whole picture of the importance of Justicia in hal-
lucinogenic preparations is available.

Chemical studies of Justicia are still incipient. Reports of the
presence of tryptamines need corroboration (Schules, R.E.:
Ann. Rev. PI.Physiol. 21(1970) 571-594: Schultes, R.E. and A.
Hofmann, The Botany and Chemistry of Hallucinogens (1973)
192).

Justicia Schultesii Leonard in Contrib. U. S. Nat. Herb. 31
(1951) 54S.

CoLomBIA: Comisaria del Vaupes, Rio Kananar, Cerro Isibukuri. August 4,
1951. R.E. Schultes et I. Cabrera 13222.

Justicia Schultesii is used by the Makuna Indians in the same
way and for the same purposes as J. Cabrerae.

Mendoncia aspera (Ruiz et Pav.) Nees in DeCandolle Prodr. 11
(1847) 51.
CoLomBia: Comisaria del Vaupes, Rio Kananar1, Cerro Isibukuri. August 3,

1951. R.E. Schultes et 1. Cabrera 13259. — Rio Apaporis, Soratama. August
24, 1951. R.E. Schultes et 1. Cabrera 13710.

The root of Mendoncia aspera is employed, crushed and
thrown into still water as a fish poison by the Taiwano Indians
(Schultes, R.E.: Lloydia 29(1966)302).

Ruellia colorata Bail/on in Bull. Mens. Soc. Linn. Paris 2 (1890)
853.
CoL_oMBIA: Comisaria del Putumayo, Rio Sucumbios, Quebrada Conejo.

‘‘Large bush. Bracts fiery red, flowers red-orange. Used as ornament in
nose, ears, wrists.’ April 2-5, 1942. R.E. Schultes 3507.

The Kofan Indians employ the root of Ruellia colorata as a
vermifuge and vomitive. The roots are pounded into a paste
which is simply kneaded in cold water. The Kofan name is
see-tu-ku-t-he-ta. In the Brazilian Amazon, the root of Ruellia
geminiflora HBK. is considered a vomitive (LeCointe, P.: A
Amazonia Brasileira 3(1934)205).

271

Sanchezia thinophila Leonard in Bot. Mus. Leafl. Harvard
Univ. 16(1953)94.

COLOMBIA: Comisaria del Amazonas, Trapecio Amazonico, Rio Loreto-
yacu. October 20-30, 1945. R.E. Schultes 6607. — Puerto Narino, near mouth

of Rio Loretoyacu. September 13-15, 1966. R.E. Schultes, R.F. Raffauf et D.
Soejarto 24121.

The Tikuna Indians of the Trapecio Amazo6nico prepare a
decoction of the inflorescences of Sanchezia thinophila which
is used as a wash to bathe the heads of girls who undergo the
ritual adolescent initiating ceremony so characteristic of this
tribe. The hair is forcefully pulled out, leading to profuse bleed-
ing.

It is possible that this use is related to a kind of ** Doctrine of
Signatures’’ connection with the large, showy, blood-red
bracts of the plant.

Little is known of the chemistry of Sanchezia. No biody-
namic constituents have been reported for the genus, but may
have a high tannin content. The leaves of Schultes, Raffauf et
Soejarto 24121 were alkaloid-negative with Dragendorff rea-
gent.

Teliostachya lanceolata Nees var. crispa Nees in Martius, FI.
Bras. 9(1847)72.

PERU: Departamento de Loreto, Iquitos and vincinity. ““Toé negra. Culti-
vated. Flowers white with violet. 0.5 - 1 m. in height.’ July 15, 1967. R.
Martin, T. Plowman et C. Lau - Cam 1638.

The Kokama Indians use this plant as an hallucinogen and as
an admixture in a narcotis preparation. Teliostachya seems not
to have been phytochemically investigated.

PLATE: 2)

i. ,
‘4

JUSTICIA

pectoralis Jacq.

Plate 21. Justicia pectoralis Jacq.

PA

PLATE 22

Plate 22. Justicia pectoralis var. stenophylla Leonard.

274

PLATE 23

=

~

MENDONCIA >

aspera

(R.&P) Nees

1

Plate 23. Mendoncia aspera (Ruiz et Pav.) Nees.

275

BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY

CAMBRIDGE, MASSACHUSETTS, NOVEMBER-DECEMBER 1978 VOL. 26, No. 9-10

A NEGLECTED MAYAN GALACTAGOGUE
IXBUT (EUPHORBIA LANCIFOLIA)

FREDERIC ROSENGARTEN, JR.*

A potentially important medicinal plant — indigenous to
Guatemala, Belice and Honduras — is the perennial herb
Euphorbia lancifolia Schlecht., commonly known as “out
in Guatemala and El Salvador, and less well known as “‘hierba
lechera’’ in southern Mexico. It is a member of the spurge
family (Euphorbiaceae).

According to Black’s Medical Dictionary: *‘Galactagogues
are drugs which increase the flow of milk in nursing women.
The normal stimulus of an infant’s lips is the most powerful
agent in producing milk, and a mother who has little or no milk
should nevertheless hold the infant to the breast. Good food
and the hormone, prolactin, from the pituitary gland increase
the quantity and improve the quality of milk.’ ’(1)

Traditional Guatemalan folklore claims that ixbut (rhymes
with ‘‘wish-boot’’), taken as an herbal tea, will stimulate lacta-
tion and increase the flow of mother’s milk in postpartum
women. A modern report in the Flora of Guatemala (2) states:
‘It is said to double the quantity of milk given by cows that eat
it. An infusion or decoction of it often is given to nursing
women to increase their flow of milk, and it is claimed that it
will cause the milk to flow after it has ceased normally, or even
in women who have not given birth to a child.”’

In this connection, one still hears countless tales about the
* Associate in Economic Botany.

Botanical Museum Leaflets ( USPS 404-990). Published monthly except during July and August by
the Botanical Museum, Harvard University, Cambridge, Massachusetts 02138. Subscription:

$25.00 a year, net, postpaid. Orders should be directed to Secretary of Publications at the above
address. Second-Class Postage Paid at Boston, Massachusetts.

wondrous powers of ixbut as a galactagogue, some of which
border on the sensational: for example, there are numerous
claims that aged grandmothers, or even great-grandmothers,
after taking ixbut, have been able to suckle newlyborn infants
through their withered breasts when the young mothers died in
child birth.

An even more curious incident involving ixbut was reported
in a Guatemalan newspaper in November 1952 (3): during the
late 1890's, a Guatemalan physician, Dr. Pedro Molina F., was
at his home near Flores, Peten. One afternoon, he received a
message that he was urgently needed by a woman in labor. By
the time he arrived at the isolated, humble, native hut, he
managed to save the life of the baby girl, but the mother died.
Dr. Molina thereupon asked the feeble great-grandfather, who
appeared to be at least ninety years old, what woman was going
to nurse the infant. This venerable progenitor replied that no
woman was around, but no woman was in fact needed since he
himself would be the wet nurse; he was going to drink a tea of
the medicinal herb ixbut which would enable him to provide
milk for his new great-granddaughter. The physician objected
and reluctantly departed. Six days later, Dr. Molina returned to
check on the condition of the baby: he found the old man
boiling ixbut leaves in a pot of water: for five days he had been
drinking the infusion, but he complained that his swollen
breasts hurt him when the infant suckled. The physician exam-
ined the great-grandfather’s breasts which indeed were
enlarged like the teats of a perfect wet nurse and were exuding a
milky juice that tasted like mother’s milk. The baby was thriv-
ing.

I know of one authentic case: in 1963, Senorita Bertha
Garcia, a teacher in the dietary and anthropological service of
the Institute of Nutrition of Central America and Panama (IN-
CAP), was on a dietary survey with an INCAP technical,
nutritional team in the village of Santa Cruz Balanya, Depart-
amento de Chimaltanango, when she met a 45-year-old Indian
woman who was nursing a small 14-month-old baby. The wom-
an’s sister had recently died in child birth, and since the family
was so poor they couldn't afford to buy milk or any other food
for the infant — this woman, who had one son of twenty-five,

278

and had not nursed a baby for a quarter of a century, took it
upon herself to nurse her infant nephew. So she started to take
a tea made from ixbut leaves, three times a day — and took it
for several months. Senorita Garcia was skeptical: upon re-
quest, the mature, nursing aunt exposed her breast, which
indeed contained an abundant supply of milk.

The word ‘“‘ixbut’’ had its origin in the early Mayan lan-
guages of the Pokom group (4) — then gradually spread to the
quiche and mam tongues of Guatemala (5). ‘‘Ix’’ meant wom-
an; while “‘but’’ stood for an increase in the flow or volume of
water. Hence the implication that the plant ixbut increased the
volume of liquid (i.e., milk) in women (4).

Euphorbia lancifolia Sch/echtendal in Linnaea 7 (1832) 143.
This herb was described by D. F. L. von Schlechtendal on
the basis of material collected in a shady forest at Misantla,
Mexico, approximately 60 miles northwest of Vera Cruz, in
early 1829, by the German collector Christian Julius Wilhelm

Schiede.
The original description by Schlechtendal follows:

36. E. lancifolia n. sp.; herbacea carnosula glabra alternifolia,
inflorescentia dichotoma terminali. Caulis articulatus, articulis
longitudine eximie diversis, glaberrimus, laevis, in statu sicco
striis elevatis notatus teres. Folia alterna breviter petiolata late
lanceolata aut rhombeo-lanceolata, basi acute apice vero acutis-
sime acuminata, superficie obscurius viridi glabra, pagina infera
dilutiori glaucescente pilisque minutis albis adpressis oculo ar-
mato tantum in conspectum venientibus adspersa, integerrima,
margine paululum inflexo, majora 3'4—3%4 p. longa, 17—19 1. in
medio lata, petiolo 4—S lineari. Flores in apicibus caulis ter-
minalem et ex ultimis axillis axillares cymas efformant
dichotomas, paucifloras abortu florum v. ramulorum lateralium,
inferiori in sua parte bracteis parvis lanceolatis acutis 1% lin.
longis oppositis viduis saepe obsessas. Involucrum campanula-
to-turbinatum 4-fidum, laciniis exterioribus inaequalibus (3
majoribus quarta) erectis (an semper?) semirotundis inferne
bilabiatis, labiis angustis lacinia brevioribus, laciniis interioribus
alternis minoribus tenerioribus apice laciniatis. Stamina s. flores
masculi more generis et flos centralis pedicellatus, dein elon-
gatus nutans, ovario dense breviter piloso. Fructus non vidimus
nec juniores. — In sylvis umbrosis Misantlae. Mart.

A contemporary description, published by Standley and
Steyermark in Flora of Guatemala (2) follows:

279

**A perennial herb, somewhat fleshy and succulent, the stems
terete, pale greenish, glabrous or nearly so, ascending or pro-
cumbent or prostrate, sometimes greatly elongate, as much as 2
meters long or more, and subscandent; leaves alternate, on very
short, stout petioles, mostly rhombic-lanceolate and 5-9 cm.
long, acute or acuminate, acute at the base, entire, green and
glabrous above, pale beneath and inconspicuously and sparsely
pilosulous or glabrate, the lateral nerves obsolete; involucres in
small, almost naked, terminal cymes, campanulate-turbinate,
4-lobate, glabrous, the lobes obovate, fimbriate, the glands
transverse-ovate, the appendage semiorbicular, crenulate, white
or whitish.”

The range of Euphorbia lancifolia is now known to include
southeastern Mexico, Guatemala, Belice, El Salvador and
Honduras. It seems to be most abundantly represented in
Guatemala where it has been recorded from Alta Verapaz,
Escuintla, Peten, Izabal, Santa Rosa, Suchitepequez, Re-
talhuleu, Guatemala, Sacatepequez, Quezaltenango, San Mar-
cos, Quiche and Huehuetenango. It prefers damp thickets,
occasionally growing, however, in pine forests or in open
fields. Ixbut may be found at many different elevations: at
1,000 feet or less on the warm lowland areas of the Pacific coast
in the Departments of Escuintla, Suchitepequez and Re-
talhuleu — to areas up to 5,800 feet and sometimes higher in the
Departments of Quezaltenango, San Marcos and the Alta Ver-
apaz, in the cool highlands.

The vernacular names of Euphorbia lancifolia have been
reported as ixbut and sapillo (2), but the plant is usually known
throughout Guatemala, and even beyond the borders of the
country, as ixbut. It was introduced to Cuba, where the name
has been recorded as isbut.

Ixbut may have originated in the warm, humid drainage
basin of the region known today as the Departamento del
Peten, Guatemala — possibly near San Benito, southwest of
Lake Petén Itza (6). (See map, Plate 28)

Ixbut plants may be readily multiplied by root cuttings. In
the Coban region (Alta Verapaz), ixbut is often cultivated as a
medicinal herb near villages, so it will be readily available as a
galactagogue.

Standley and Steyermark (2) report: *‘Rather curiously, it is
claimed in Coban that the plants often cause the death of cattle

280

and horses eating them, and this may be the result of the
inherent properties of the seeds.”’

I have never heard reports during my field work in
Guatemala that would indicate toxicity in Euphorbia lan-
cifolia; nor in my discussions with local physicians has any
such danger been expressed. It should be emphasized that the
leaves and branchlets of ixbut, not the seeds, should be
utilized. Furthermore, the seed, which is minute, apparently is
not used by the native population. It is probable that the report
about the death of horses and cattle is the result of a confusion
of ixbut with another plant, possibly another species of
Euphorbia, which genus does have toxic species.

Roys points out (7) that in ancient Maya medicine, in the
‘‘Ritual of the Bacabs,’’ the Maya doctors believed in curing
like with like: a vine that looked like a serpent was considered a
fitting cure for snake bites; a yellow fruit was given as medicine
to cure jaundice: red plants or fruits were prescribed for pa-
tients who vomited blood or suffered from dysentery. Since
ixbut contained an abundant, sticky, milky sap, which exuded
from its broken stems and leaves, it was fitting that it may have
been recommended to increase milk flow in nursing mothers.

Erwin P. Dieseldorff, a German naturalist who settled in the
Coban region of Guatemala during the latter part of the nine-
teenth century, included ixbut in a booklet which he wrote on
medicinal plants of the Alta Verapaz (10). He described it as a
prevalent herb growing especially well in damp areas. The dark
green ixbut leaves, containing a milky latex, should be cooked
in water together with their tender branchlets to prepare an
herbal tea, to be used as a galactagogue.

In March 1971, the Instituto Indigenista Nacional de
Guatemala published a bulletin (9) concerning popular
medicine in rural areas of Guatemala. The medicinal plant
ixbut is mentioned several times in this report, especially to
increase lactation in postpartum nursing mothers. It was listed
as a popular folk remedy in Coban and Chamelco, in the zone of
Kekchi; and in San Cristobal Verapaz and Tamahu in the zone
of Pocomchi.

Usually it is recommended that the ixbut leaves be boiled in
water: the liquid allowed to cool; sugar added, and the drink

281

consumed three times a day. In Tamahu, it is likewise sug-
gested that fresh ixbut leaves be eaten, uncooked, like a salad.

In another portion of this same report, fresh ixbut leaves are
recommended as a cure for sexual impotence, in the village of
San Cristobal Verapaz.

By 1945, Euphorbia lancifolia had been introduced to Cuba
(8), where it was recommended as a galactagogue. Just as in
Guatemala, it was reported in Cuba that ‘‘isbut’’ could increase
lactation up to 100% in nursing mothers.

In 1949, Dr. Manuel Serrano, a botanist, chemist and pioneer
in the serious testing and utilization of ixbut in Guatemala,
reported ona series of controlled tests that had been carried out
ona group of nursing mothers at or near the General Hospital in
Guatemala City — in collaboration with Merck & Co., Inc.
(11). Of the 86 postpartum women tested, 54 showed an abun-
dant increase in the production of milk after taking ixbut. In
some cases, mothers who during earlier parturitions had been
unable to nurse, were able to do so effectively after drinking an
herbal tea of ixbut for several days. In one case, following a
Caesarean section, the milk secretion had been almost
nonexistent: 48 hours after the administration of ixbut, how-
ever, the lactic secretion increased to such an extent that not
only was the child thoroughly satisfied, but the excess of
mother’s milk soaked the sheets during the night.

The results of the ixbut tests were carefully checked by Dr.
Serrano. The duration of the therapy was usually from 3 to 5
days. The parts of the ixbut plant utilized were those above the
ground, i.e., the stems, leaves and flowers — but not the roots.
The average dose of ixbut given to the women: 5 leaves or 5
sections of stem (about 5 grams) were brewed to make a cup of
herbal tea; then 6 cups of ixbut tea were given daily to each
patient.

The chemical analysis of the mothers’ milk carried out be-
fore and after the intake of ixbut showed that the quality of the
milk did not undergo any notable change in its composition,
even though the ixbut may have excited additional secretion.

Dr. Serrano cautioned that the ixbut tea should not be al-
lowed to ferment: when it turns yellow, it should be discarded
since it might cause diarrhea.

282

Time magazine published an article in 1949 (12) entitled
‘‘Milkweed,”’ in a section devoted to Medicine. It described
some of Dr. Manuel Serrano’s experiments with ixbut in
Guatemala: controlled testing of 1,800 women who had experi-
enced trouble in nursing their babies indicated that 50% could
not nurse at all without ixbut; 35% who could nurse only a little
showed notable improvement after taking ixbut; while 15% did
not benefit from ixbut.

Dr. Serrano pointed out to Time that he was convinced that
ixbut increased milk flow — actually milk, and not water. The
active principle in ixbut that caused the increase in lactation
had not been isolated in 1949. (Today, thirty years later, it still
has not been determined just what this active principle may
be.) Dr. Serrano in 1949 ruled out the milk-producing hormone
prolactin: while prolactin did not enable his wife to nurse her
four babies, ixbut did.

Between 1949 and 1951, the late Dr. Efren C. del Pozo of
Mexico City, former Secretary General of the Union de Uni-
versidades de America Latina, carried out considerable re-
search on ixbut in collaboration with Merck & Co., Inc., both
in Mexico City and in Guatemala City (13).

During 1949, Dr. del Pozo’s first testing was made at the
Centro Materno Infantil Avila Camacho in Mexico City on 21
nursing women. The ixbut plant material had been shipped by
Air Cargo from Guatemala City and was thus from 4 to 7 days
old. During 1950, Dr. del Pozo carried out further research at
the Mexico City Penitentiary: 14 lactating mothers were tested:
11 were given ixbut, while 3 were left as controls. In 1951, a
third series of tests involving ixbut took place in Guatemala
City under Dr. del Pozo’s direction, at the Maternidad de
Guarderias Infantiles, on 22 nursing women and 3 non-lactating
women.

On the whole, the results of Dr. del Pozo’s experiments on
the effect of ixbut were inconclusive and somewhat negative,
even though in some nursing mothers a variable increase of
lactation was noted following treatment with an aqueous
extract of ixbut; average dose approximately 750 cc., a 10%
infusion, daily for 25 days.

Dr. del Pozo made the following observations following
three years of research on 1xbut:
(1) For an accurate evaluation of any galactagogue such as
ixbut, a long control period is necessary, since short term trials
are unreliable;
(2) Asaresult of his studies, Dr. del Pozo concluded that ixbut
does not induce lactation in non-lactating women — despite
popular, folkloric claims in Guatemala to the contrary.
(3) Ixbut does not produce breast pain or congestion in lacta-
ting women between the second and fourteenth month of lacta-
tion;
(4) Ixbut does not produce any appreciable change in breast
volume or in the mammary glands of nursing mothers;
(5) The active principle (if any) of ixbut does not seem to be
unstable. Very fresh plant material administered in Guatemala
did not show higher activity than material several days old.
This fact is also contrary to popular belief in Guatemala;
(6) Ixbut seems to have higher activity as a galactagogue in
Guatemala during the month of November, i.e., in the early
part of the dry season in Guatemala City shortly before the
plant blossoms;
(7) On the whole, in the opinion of Dr. del Pozo, the lactogenic
properties of ixbut have been considerably exaggerated — not
only in the case of women, but also when he carried out ixbut
tests on postpartum female cats:
(8) Dr. del Pozo, however, pointed out that the striking effects
of ixbut reported in Guatemala by lay people cannot be totally
denied on the basis of his testings. The Guatemalan claims refer
mainly to ixbut used during the first days after delivery when
lactation is started; while his tests were designed usually to
explore lactagogue effects from two to fourteen months after
childbirth;
(9) Furthermore, Dr. del Pozo personally knew of the case ofa
woman in Mexico City who had to feed artificially two of her
previous children for lack of natural milk. When she received
infusions of fresh ixbut, she was able to breastfeed her third
child with a remarkable increase of the volume of milk.
During 1952, Merck & Co., Inc. discontinued its research on
ixbut. It was felt that although ixbut may have some effects as a

284

galactagogue during the first days of lactation, these results
were highly irregular and difficult to prove.

Ixbut has been used, off and on, for many years to supple-
ment cattle feed in Guatemala (4). In 1894, Fermin Rosal noted
that his own cows in Cuyotenengo, on the Pacific coast of
Guatemala, were not producing anywhere nearly as much milk
as the cows on the neighboring hacienda which were being fed
ixbut.

In 1911, Eduardo Saravia Castillo elaborated a Guatemalan
product called “‘“GALAC-LATEX”’ as a supplement for cattle
feed, in which ixbut was the principal ingredient. This galac-
tagogue was apparently successful for a few years, but it is no
longer produced.

In June 1927, Professor Guillermo Gandara carried out an
experiment in La Ceiba, El Salvador, concerning the effect of
ixbut on milk production in cows (14). The poorest producer
was selected from 8 milch cows. This cow had been producing
on the average 2 bottles of milk (each bottle roughly 3/4 liter)
daily. For 5 consecutive days the cow was given, as a dietary
supplement, an infusion of ixbut — 250 grams per liter of water;
then normal forage for 3 days; then 3 days more of the dietary
supplement of ixbut. At the end of 11 days, this cow had tripled
its production of milk from 2 bottles to 6 bottles per day.

In 1947, J. Ignacio Aguilar noted in southeastern Mexico that
ixbut would increase milk yields in cows, when mixed with
other cattle forage (15). He recommended mixing ixbut in small
quantities — about 1.5 grams ixbut leaves per kilo of the cow's
weight — in an aqueous solution to supplement other cattle
feed in order to provide a balanced diet for dairy cows. He
observed, however, that too much ixbut should not be fed to
the cows: for a few days milk yields would increase substan-
tially, but then gradually decrease as the animals’ physical
condition deteriorated, possibly due to overstrain and exhaus-
tion.

According to Aguilar, ixbut is a galactagogue par excellence
for dairy cows, superior to alfalfa and other tropical, legumin-
ous forage plants. Ixbut is extremely vigorous and more resis-
tant to plant diseases than alfalfa. For best results, ixbut should
be planted in warm, shady, protected areas where there is

285

constant humidity. It is readily propagated by root cuttings; it
should be planted in rows 35 centimeters apart, with approxi-
mately 25 centimeters between the plants in the rows. After
two years, under favorable conditions, an average of 25 tons of
green ixbut forage may be harvested per hectare per year; with
two cuttings in the rainy season and one in the dry season. Each
time the forage is cut, new growth is stimulated. If cared for
properly, an ixbut pasture should last 15 to 20 years.

Ina laboratory analysis, Aguilar (15) found that ixbut leaves
(Euphorbia lancifolia) contained 7.46 per cent protein, as com-
pared to 4.72 per cent for alfalfa (Medicago sativa).

In 1949, Dr. Manuel Serrano (11), in collaboration with
Merck & Co., Inc., investigated the effect of ixbut on the milk
production of 6 goats in Guatemala City. The goats were given
a 5% aqueous infusion of fresh ixbut plant material daily, fora
period of 4 days. No signs of toxicity were noted. A modest
increase in goat’s milk production was observed in each case.

During 1949, tests were made of the insecticidal properties of
78 species of plants from tropical America used as fish poisons,
insecticides or drugs (16). Through several years, roots, stems,
leaves, flowers and seeds were collected and tested at the
Puerto Rico Agricultural Experiment Station, Mayaguez,
Puerto Rico, to determine the toxicity of the materials to in-
sects.

Five species were found to be highly toxic; 30 species par-
tially toxic; while 43 species, including ixbut (Euphorbia lan-
cifolia) were found to be non-toxic to insects.

In 1957 (17), Squibb, Braham and Scrimshaw carried out
experiments on the utilization of the carotenoids of ixbut
(Euphorbia lancifolia), teosinte (Euchlaena mexicana), yellow
bamboo leaves (Bambusa vulgaris) and green bamboo leaves
(Bambusa ventricosa), by New Hampshire chicks over a
5-week feeding period. The carotene content and vitamin A
activity of dehydrated forage meals prepared from these in-
gredients were studied. It was found that ixbut contained the
highest carotene content of the four forages and maintained the
highest serum levels of vitamin A. The dehydrated meals pre-
vented mortality and permitted good weight gains.

286

The chemical composition of these dehydrated forage meals
was as follows:

Yellow Green

Ingredients Ixbut bamboo bamboo Teosinte
leaves leaves

Ash, gm.% 9.8 21.3 19.7 12.5
Calcium, mg.% 1,519 - — 1,163
Iron, mg.% Sa.5> ios 8.0 147.6
Phosphorus, mg.% 400 86 86 320
Moisture, gm.% 9.2 8.6 6.6 4.0
Ether extract, gm.% 5.9 Zio 2:3 331
Crude fiber, gm.% 162 21.7 24.4 21.6
Nitrogen, gm.% 1.90 1.61 1.55 1.20
Carotene, mg.% 15.05 12:32 8.61 8.28
Vitamin C, mg.% 73 6 8 38
Vitamin Bz, mg.% 0.66 0.90 0.65 0.57
Vitamin Bi, mg.% 0.04 0.10 0.16 0.10
Niacin, mg.% 4.22 2.54 3.10 3.32

The following chromatographic analyses of dried 1xbut
leaves from Guatemala City, Guatemala, were reported by the
chemical research laboratory of SISA Incorporated of Cam-
bridge, Massachusetts, on August 21, 1979:

EXTRACTION OF EUPHORBIA LANCIFOLIA (IXBUT)

In order partially to characterize some of the principles of
Euphorbia lancifolia, an extract of dried ixbut leaves was
prepared and the extract subjected to gas-liquid chromatogra-
phy (figure 1) and high pressure liquid chromatography (figure
2).

The gas-liquid chromatographic analytical method separates
the components of the extract primarily on the basis of their
volatility with the most volatile materials appearing first.

The high pressure liquid chromatographic analytical method
separates the components primarily on the basis of polarity.
With a ‘‘reverse phase’ column of the type used here, the more
polar components appear early on the chart.

Since in both of these methods each peak on the chart corre-

287

sponds to at least one component in the mixture, the results
shown in figure | and figure 2 indicate that the extract contains
a large number of components. These components have not
been identified but the results indicate that these methods,
particularly the high pressure liquid chromatographic method,
could be used to separate some of the components in quantities
large enough for further chemical and biological characteriza-
tion. The analytical scans are, in effect, the signature of ixbut in
chromatography under the stated conditions, for future refer-
ence.

PROCEDURE

A 5.0 g sample of dried leaves of Euphorbia lancifolia was
homogenized briefly with 100 ml of ethanol in a blender. The
mixture was transferred to a Soxlet apparatus and extracted at
reflux with 250 ml of 95% ethanol. These extraction conditions
were chosen to approximate the use of the plant by boiling the
leaves in water to give an herbal tea.

The ethanolic extract was subjected to gas liquid (glc) and
high pressure liquid (hplc) chromatographic analyses.

The glc analysis was carried out on a 6ft x 1/8 inch stainless
steel column of 2% OV-17 on 100/120 mesh Supelcoport using a
Varian 1400 instrument equipped with a temperature pro-
grammed run from 100°C to 340°C at 20°/min. temperature rise.
The gle scan for this run is shown in figure 1.

The hple analysis was done on a 30 cm x 3.9 mm “‘reverse
phase’ column of « Bondapak Cis using a Water ALC 200
instrument with an ultraviolet (254 nm) detector. A satisfactory
analysis was obtained using acetonitrile as the mobile phase.
The hple scan for this run is shown in figure 2.

According to Dr. Juan Jose Urrutia, Chief, Nutrition and
Infection program of the INCAP, ixbut is rarely used at the
present time as a galactagogue in Guatemala City. On the
whole, the pure Indian in the rural areas of Guatemala pro-
duces more milk for breast feeding than the urbane mother of
white or mixed racial background in the city.

288

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In remote regions of Guatemala, the Indian women today
still utilize ixbut, but only when they have trouble producing
breast milk, especially during the first 3 days postpartum. In
these far-off localities, the baby has no other food or water, and
depends on his mother’s milk for both, during a period of 2
months or longer.

In Guatemala City, on the other hand, the urbane women are
no longer interested in ixbut; they can obtain other foods for
the baby such as powdered milk, bottled milk, eggs and veget-
ables.

Dr. Juan José Hurtado V., a specialist in pediatric medicine
in Guatemala City, concurs that ixbut is used much less in
Guatemala in 1979 than it was 30 years ago when he was a
resident intern. In those days, Dr. Hurtado heard a great deal
about the merits of ixbut as a galactagogue: midwives fre-
quently recommended it. In fact, when Dr. Hurtado was born
in Guatemala City in 1926, his mother took an infusion of ixbut
leaves three times a day for several days and was able to
breast-feed him. At that time, ixbut was a ‘‘fashionable”’
medicinal herb, readily available in several local markets. At
the present time, it is vitually impossible to find ixbut in any
market in or near Guatemala City. According to Dr. Manuel
Serrano, ixbut is still sold in small quantities, from time to time,
in the more remote, indigenous markets of San Felipe Re-
talhuleu, Coban and Colomba.

Dr. Hurtado categorizes ixbut as a non-toxic, beneficial,
medicinal herb, rarely used or even mentioned today in
Guatemala City.

The American Academy of Pediatrics and the Canadian
Pediatric Society have recently issued a joint statement to
encourage all physicians to recommend breast-feeding (18).
The ‘‘breast is best’? movement is apparently gaining in popu-
larity. The Harvard Medical School Health Letter stresses the
critical role of breast feeding in preventing gastroenteritis in
infants in developing countries. Respiratory infections, men-
ingitis and other overwhelming infections are less frequent
among breast-fed infants.

Mrs. Dina Nathusius is one of the Guatemalan leaders of La
Leche League International, Inc., of Franklin Park, Illinois —

291

a group interested in fomenting natural nursing of babies
through breast-feeding. Mrs. Nathusius points out that in the
local Guatemalan chapter of La Leche League, young mothers
are advised it is perfectly allright to take ixbut if they want to,
just as their mothers and grandmothers used to many years
ago, since it is not toxic and might indeed help them psycholog-
ically — by giving them confidence, by relaxing them and so
helping them to nurse their babies. Some young mothers have
indeed informed Mrs. Nathusius that ixbut has aided them to
breast-feed their babies. Mrs. Nathusiu confirms, however,
that very little ixbut is utilized at the present time in Guatemala
in comparison to two generations ago.

In Maternity Nursing (19), it is emphasized that worry and
emotional tension have an adverse effect on lactation in the
nursing mother. Perhaps, as Mrs. Nathusius suggests above,
ixbut itself may at times be beneficial as a tranquilizer — due to
its soothing, relaxing effect upon those postpartum mothers
who have confidence in its power as a tried and true galac-
tagogue.

In 1950 (13), Dr. Efren del Pozo, in fact, found in experi-
ments of his own as a Mexican physician, that intravenous
injections of ixbut caused a fall of blood pressure and a marked
slowing of the heart, in tests carried out on lactating women in
the Mexico City Penitentiary.

Dr. Lewis H. Sarett, Senior Vice President, Science and
Technology, of Merck & Co., Inc., has the following com-
ments to make about ixbut: to date the active principle of this
galactagogue has not been isolated: if found, it would consist
probably of pure crystals resulting from a chemical fractiona-
tion of the fresh plant material.

Itis a long and costly process now to introduce a new drug.
The problems following Thalidomide during the late 1950’s and
early 1960's led to the Kefauver-Harris amendments of the
Pure Food and Drug laws in 1962 — designed to prevent a
recurrence of the Thalidomide tragedy in any new drug.

Although there is a trend towards a greater utilization of
products of nature, there are much longer delays in gaining
Food and Drug Administration approval for their use. For
example, even if ixbut, let us say, should prove to be an

fis Ps

effective galactagogue for dairy cows, the F. D. A. might take a
dim view of adding a drug orally to cattle feed which would
show up in milk for human consumption — unless it were
completely metabolized in the liver or in the intestinal tract.

In order to gain approval of drug regulatory agencies in most
countries, it would be necessary to measure the level of the
product (ixbut) and its metabolites in milk. An adequate safety
ratio would have to be determined: all major metabolic prod-
ucts should be checked to determine their toxicoligical effects
on at least two species of animals, with the drug adminstered
over a period of several years. Extreme caution would be
necessary.

CONCLUSION

Ixbut, Euphorbia lancifolia, is a potentially important, natu-
ral galactagogue of Central American origin, worthy of further
chemical and pharmacological study.

This medicinal herb has been used for centuries in
Guatemala, in Maya medicine, to increase the flow of milk in
postpartum, lactating mothers. If used in moderation, it seems
to have no toxic side effects in humans.

Ixbut, when mixed with cattle fodder, is reputed to have
increased milk yields in cows.

Since the active principle of ixbut has never been isolated,
considerable research would be necessary before any valid
conclusions could be drawn concerning its future importance
in medicine or in the dairy industry.

The voucher collection of leaves of Euphorbia lancifolia
upon which this article is based (Frederic Rosengarten, Jr. s.
n., July 10, 1979, Guatemala City, Guatemala) has been depo-
sited in the Botanical Museum of Harvard University.

ACKNOWLEDGMENTS

I wish gratefully to acknowledge the valued cooperation and
orientation of Dr. Manuel Serrano, Universidades

293

Guatemala-Alemania, Quezaltenango, Guatemala, who
provided me with much pertinent information concerning the
folklore and other ethnobotanical data available on ixbut, as
well as several relevant photographs.

I also wish to express my appreciation to the following
persons who have provided assistance to me during the prepa-
ration of this article: to Dr. Richard Evans Schultes for his
thoughtful editorial guidance; to Dr. Lewis J. Sarett and Mrs.
Martha Windholz of Merck & Co., Inc., Rahway, New Jersey;
to the late Dr. Efren C. del Pozo of Mexico City; to Dr. Elfriede
de Poll, Lic. Mario Dary Rivera and Prof. Jose Esteban
Jimenez Ochoa of the Botanical Garden, Guatemala City; to
Dr. Juan Jose Hurtado V. of Guatemala City; to Dr. Juan José
Urrutia and Senorita Bertha Garcia of the INCAP, Guatemala
City; to Dr. Norman R. Farnswoth of the University of Illinois
at the Medical Center, Chicago; and to Mr. Franz J. Ippisch,
Mrs. George-Ann Mellen and Mrs. Dina Nathusius of
Guatemala City.

ILLUSTRATIONS: Plates 24, 25, 27, 30 and 31 by A. Gutt;
plate 26 from Las Plantas Medicinales del Departamento de
Alta Verapaz, by Erwin P. Dieseldorff, 1940; plates 28 and 29
by Elmer W. Smith; plates 32, 33, 34 and 35 by Dr. Manuel
Serrano.

LITERATURE CITED

_—~

1) Black’s Medical Dictionary. 1974. Thirtieth Edition. Barnes and Noble
Books. New York.

2) Standley, Paul C. and Steyermark, Julian A. 1949. Flora of Guatemala.
Fieldiana: Botany. Vol. 24, Part VI, 106. Chicago Natural History
Museum.

( 3) Ellmparcial, Guatemala. November 21, 1952. **La Planta lechera,”’ por

el Dr. Olindo Secondini B.

( 4) Serrano, Manuel. 1978. Informe cientifico sobre la planta Euphorbia

lancifolia. Quezaltenango, Guatemala. (Unpublished)

( 5) Instituto Indigenista Nacional, No. 10. 1950. Alfabeto para los cuatro

idiomas indigenistas mayoristas de Guatemala. Guatemala.

( 6) Exposicion Nacional. Fiesta de Minerva. 1913. Catdlogo de Plantas

Reputadas Medicinales en la Republica de Guatemala, 40.
( 7) Roys, Ralph L. 1931. The Ethno-Botany of the Maya. Middle American

294

_

(18)

(19)

Research Series, No. 2. The Tulane University of Louisiana, New
Orleans.

Roig y Mesa, Juan Tomas. 1945. Plantas Medicinales, Aromaticas y
Venenosas de Cuba. Parte I. Habana. 384.

Guatemala Indigena. Marzo de 1971. Aspectos de la Medicina Popular
en el Area Rural de Guatemala. Vol. VI, No. 1.

Dieseldorff, Erwin P. 1940. Las Plantas Medicinales del Departamento
de Alta Verapaz. Second Edition. Guatemala. 1977, 45.

Serrano, Manuel. 1949. ‘‘Ixbut, Euphorbia lancifolia Schlecht. en la
Republica de Guatemala.’’ Unpublished report for Merck & Co., Inc.

Time magazine. August 1, 1949. 38.

del Pozo, Efrén C. 1949-1952. Unpublished reports concerning ixbut for
Merck & Co., Inc.

Gandara, Guillermo. 1929-1930. *‘La Hierba de la Leche, Euphorbia
lancifolia..’ Memorias y Revista de la Sociedad Cientifica Antonio
Alzate. Tomo 52. Nos. 5-8, 237-246. Mexico.

Aguilar, G., José Ignacio. 1947. Forrajera de porvenir para la ganade-
ria. Ixbut. El Rancho Mexicano. 3 (24). 8-9.

Sievers, A. F., Archer, W. Andrew, Moore, Rufus H. and McGovran,
E.R. 1949. Insecticidal Tests of Plants from Tropical America. Scien-
tific Notes. Journal of Economic Entomology. Vol. 42, No. 3, 549-551.

Squibb, Robert L., Braham, J. Edgar and Scrimshaw, Nevin S. 1957.
Utilization of the Carotenoids of Bamboo Leaves, Teosinte and Ixbut
by New Hampshire Chicks. Poultry Science. Vol. 36, 1241-1244.
Guelph, Ontario, Canada.

The Harvard Medical School Health Letter. February, 1979. Vol. IV,
No. 4. 3.

Reeder, Sharon R., Mastroianni, Luigi, Jr., Martin, Leonide L. and
Fitzpatrick, Elise. Maternity Nursing. Thirteenth edition. J. B. Lip-
pincott Company, Philadelphia. 362.

295

PLATE 24

_

=
a
_

Plate 24. The rank, much-branched, perennial herb, ixbut — 7 years old.
Botanical Garden, Guatemala City.

297

PUATE 25

Plate 25. The rhombic-lanceolate, dark green ixbut leaf, 5 to 9 cm. long, is
usually characterized when fresh by a unique white “‘V"’ , stretching from
midrib to margin.

298

PLATE 26

ITxubut

Plate 26. Drawing of ixbut from Erwin P. Dieseldorff's Las Plantas Medi-
cinales del Departamento de Alta Verapaz. Note the typical V-markings on
the leaves.

299

PLATE 27

—_
—
jeer
™
sere
—
—
| eon
soem
eee

-
—
pole
ame
~
=
ee
A »

‘
,

LAN(

Plate 27. Ixbut in the Botanical Garden, Guatemala City. The leaves con-
tain a milky latex, possibly the source of the active galactagogue principle.

300

PLATE 28

KNOWN RANGE OF IXBUT
(Euphorbia lancifolia Schlecht)

IN GUATEMALA AND ADJACENT COUNTRIES
’ Stars indicate countries or subdivisions thereof, in which
; collections have been made
Stippled area indicates locality where folk use as a galactagogue
ra may have originates ;

4

Quintana Roo
(MEXICO)

ip

—-

S--===" "1 Campeche (mexico) |
_---, Tabasco
\ MEX.
> Mex) +
\ Petén
*
Lago Feten
ei DELICE
© —Flores
sede
*
Chiapas
(MEXICO)
Ps
/, wt} wa ca \ ‘a 7
o : ae
* ri M ¥ J * of vases
( % ; * é
( * / oon yw > se}
’ _- anes nango at, | ee 7
~f x a ae & SN eT) \ ou,
rae > \ a 7 Ag™ i ¢
* yt I , org; . \
of \ U> a . : *
Quetzaitenangow ¥_-~7” 4 pe a ea A HONDURAS

301

EXPLANATION OF PLATE 29

Flowering stems, about one half natural size.
Root system with nodules, about one half natural size.
Inflorescence (cyathium), showing nectar glands, the four fimbriate lobes
of the cup, and the single, pendent, developing female flower, about
seven times natural size.
One segment of the dried dehiscent seed capsule, about six times natural
size.
Fresh seed, about six times natural size.

Botanical Illustration by E. W. Smith

302

PLATE 29

EUPHORBIA
lancifolia
Schlecht.

303

30

PLATE

—

Oy,

<
—)
—|
we
<
<—

Plate 30.

Ixbut in the Botanical Garden, Guatemala City.

304

PLATE 31

Close-up view of root system of ixbut.

Plate 31.

305

PLATE 32

Plate 32. An 18-year old, postpartum, Indian mother selects ixbut leaves for
brewing in her herbal tea. Quezaltenango, Guatemala.

306

PLATE. 33

Plate 33. The traditional Guatemalan Indian preparation of ixbut tea: brew
about 5 grams (5 fresh, macerated ixbut leaves) in 250 cc. of boiling water:
allow to cool; add sugar as desired.

307

PLATE 34

ete
i

at.

ae _. P
a » w Omwy"s
5 «~ etl * '’

Plate 34. Dosage of ixbut herbal tea, traditionally recommended in
Guatemala: 6 cups daily for 3 to 5 days to promote lactation. (Daily dose of 30
grams of ixbut in infusion.)

308

PLATE. 33

Plate 35. 24 hours after this young Indian mother had taken ixbut tea, she
was able successfully to breast-feed her baby.

309

BOTANICAL MUSEUM LEAFLETS VoL. 26, No. 9-10
NOVEMBER-DECEMBER 1978

EVOLUTION OF THE IDENTIFICATION OF THE
MAJOR SOUTH AMERICAN NARCOTIC PLANTS

RICHARD EVANS SCHULTES!,?

There are usually two events to consider in the identification
ofa narcotic: 1) the first description or reference to its use in the
literature or some other evidence — such as archaeological —
of its use: and 2) the earliest botanical collection or identifiable
illustration definitely associating the plant with its utilization as
an inebriant.

It should be then at once obvious that the basic step in our
understanding of any hallucinogen or other psychoactive plant
must necessarily rest upon a correct identification. Clearly no
chemical or pharmacological studies can be carried out on a
sound basis, until certainty concerning the vegetal source be
established: and anthropological or sociological investigations
likewise must suffer from lack of full and accurate botanical
determination of the source plant. The exact Latin binomial,
ideally backed up with a voucher specimen filed away ina
recognized botanical institution, is almost always essential to
progress in understanding any drug.

Contrary to widespread belief, the botanical identification of
medicinal. toxic and narcotic plants is not an easy task. Often
frustratingly slow, it frequently demands years of field work.
Sometimes — though fortunately not often — aboriginal users
oppose study by outsiders of their sacred plants. More fre-
quently, however, obfuscation resulting from careless, hasty
or incompetantly gathered or interpreted data by well meaning
but untrained investigators clouds the issues for many years.
There are sundry instances where, even today, we are beset
with doubts about the true and complete source of hallucino-
genic preparations.

'Delivered at the conference Hallucinogens in Native American Shamanism and
Modern Life, San Francisco, California, September 30, 1978.

2Paul C. Mangelsdorf Professor of Natural Sciences, Harvard University; Director,
Botanical Museum of Harvard University.

311

The apparently wide use of vegetal additives to alter the
effects, strength and duration of the activity of some of the
principal plant components of biodynamic drinks, snuffs and
chews is still in a very preliminary stage of investigation. It
promises to disclose a rich understanding of often hidden plant
properties on the part of the South American aborigines.

In view of the present state of our knowledge, then, it may be
advantageous to review the steps in identifying a number of the
major psychotropic plants currently used in South America
and to look at what little we know about some of the additives
employed in a few of the hallucinogenic preparations.

ERYTHROX YLACEAE

Erythroxylon* P. Brown

Coca must be rated as the most important and culturally
significant narcotic of South America. It likewise is the psy-
choactive plant that, of the New World flora, has played the
greatest role in Western medicine. Yet our understanding of the
source plant or plants has been woefully inadequate. Recent
studies are beginning finally to clarify some of the botanical and
ethnobotanical mysteries surrounding coca and its aboriginal
use.

The chewing of coca — practiced by millions of Indians in
the Andes and in the western Amazon — represents probably
the oldest use of a narcotic in South America. The oldest
archaeological records of coca-chewing date from at least 3000
B.C. Figurines of this remote date demonstrating the pouched
cheeks of coca-chewers have been discovered in sites of the
Valdivia Culture of coastal Ecuador (Lathrap, 1979). This dis-
covery represents the earliest known specimen of a continuous
series of Ecuadorian figurines showing coca-chewing, primar-
ily in the northern highland area of the country.

Actual pre-Incan coca leaves have been found ina grave site
located along the coastal area of Peru, with a radio carbon
dating of 1314 B.C. (Lanning, 1967).

*This generic epithet is often spelled Erythroxylum and it has been suggested that
Erythroxylum is nomenclaturally correct (Plowman, 1976). In this non-taxonomic
article, I prefer to employ the more widely used and more familiar Erythroxylon.

a2

The first writer to discuss coca use appears to be Monardes
in 1577. An English translation of his writing appeared in 1596
under the title Jovfull Newes out of the New-found World
(Frampton, 1596).

The first voucher botanical specimens of coca were collected
in Provincia Sica Sica in the Ungas region of Bolivia by the
French botanist Joseph de Jussieu in 1749. The species Ery-
throxylon Coca was described in 1786 by the French naturalist
Jean Baptiste Lamarck. There are three duplicate types of this
species preserved in the herbarium of the Jardin des Plantes in
Paris, together with illustrations made by Jussieu.

It has long been widely believed that there is in reality only
one species of Erythroxylon cultivated and employed as a
narcotic. As in the case of Cannabis — where many botanists
now recognize three, not one, species — recent intensive re-
search has established the fact that two cultivated species are
involved in coca use: E. Coca and E. novogranatense (Morris)
Hieron. (Plowman, per. comm.).

Although Erythroxylon Coca has been botanically known
since 1786, a precise understanding of the taxonomy of this
group of plants has, as a result of intensive field work, been
available only in very recent times due primarily to the investi-
gations of Timothy C. Plowman.

Species indicated in the foregoing discussion:

Erythroxylon Coca Lamarck, Encycl. 2 (1786) 393.
Erythroxylon novogranatense (Morris) Hieronymus in Engler, Bot. Jahrb.
20, Beibl. 49 (1895) 35.

MyYRISTICACEAE

Virola Aublet

One of the most important — yet most recently identified —
of the South American hallucinogens is that prepared from a
blood-red resin-like exudate of the inner bark of the myris-
ticaceous Virola. It is most commonly employed as a snuff and
has various names, differing according to tribe and locality.
The best known names, however, are yakee and yato in Col-

313

ombia, parica, epena or nyakwana in Brazil (Schultes, 1954a,
1954b).

Although used in the western Amazon generally, Virola as a
narcotic is most intensely utilized in the northwesternmost part
of Brazil, adjacent parts of Colombia and the Orinoco head-
water areas of Venezuela: a region where Spruce concentrated
several years of his field research, and where he devoted spe-
cial attention to the diversity of species of Virola; yet he failed
to discover this interesting native use of the genus.

It seems that the first mention of the snuff is that of the
German ethnologist Theodor Koch-Grtinberg who, in 1923,
referred to an intoxicating powder called hakudufha prepared
from a tree-bark by the Yekwana Indians of the Orinoco
(Koch-Grunberg, 1923). ‘This is a magical snuff’, he wrote,
‘*exclusively used by witch doctors and prepared from the bark
of a certain tree which, when pounded up is boiled in a small
eathenware pot, until all the water has evaporated and a sedi-
ment remains at the bottom of the pot. This sediment is toasted
in the pot over a slight fire and is then finely powdered with the
blade of a knife. Then the sorcerer blows a little of the powder
through a reed . . . into the air. Next, he snuffs, whilst, with
the same reed, he absorbs the powder into each nostril succes-
sively. The hakudufha obviously has a strongly stimulating
effect, for immediately the witch doctor begins singing and
yelling wildly, all the while pitching the upper part of his body
backwards and forwards.”

Koch-Grunberg did not authenticate this discovery with bo-
tanical material, but there would appear to be little doubt that
his description applies to Viro/a-snuff.

The earliest botanical association of an inebriating snuff with
the Myristicaceae or Nutmeg Family appeared in 1938. The
Brazilian botanist Adolpho Ducke indicated that the ‘‘ Indians
of the upper Rio Negro use the dried leaves of this species
[Virola theiodora| and of V. cuspidata in making a snuff pow-
der that they call parica’’ (Ducke, 1938). A year later, in writing
about the leguminous tree Anadenanthera peregrina, the seeds
of which are the source of yopo-snuff, he elaborated as follows:
‘*Martius and other writers attribute to this species the source
of the narcotic parica employed by certain Amazonian Indians

314

... . Notwithstanding, according to information which I ob-
tained from the natives themselves in two localities in the upper
Rio Negro, the paricd-powder comes from the leaves of
species of Virola . . ..’ (Ducke, 1939). It is now known with
certainty that the leaves are not used, but Ducke’s reports
represent apparently the earliest identification of this narcotic
powder with the correct genus of trees.

It was not until 1954, however, that the preparation of the
snuff was described in detail and specifically identified on the
basis of voucher botanical material. During my ethnobotanical
studies in Amazonian Colombia, I encountered the Indians of
the Vaupés region preparing a brownish snuff taken for cere-
monial inebriation only by witch doctors for diagnosis and
treatment of disease, prophecy and divination and for a variety
of other magical purposes. It was prepared from the inner bark
of Virola calophylla (Spr.) Warb. or V. calophylloidea Mark-
graf (Schultes, 1954a, 1954b). Later field work indicated that V.
elongata (Benth.) Warb. is likewise a source of the drug
(Prance, 1970, 1978; Brewer-Carias and Steyermark, 1976).

Gradually, studies established the fact that the most intense
use of Virola-snuff occurs amongst the large group of Indian
subtribes known collectively as the Waikas, inhabiting the
upper reaches of the Orinoco and the northern tributaries of the
Rio Negro of Venezuela and Brazil, respectively. These In-
dians refer to the snuff usually as epena, ebene or nyakwana.

The main source of the powder amongst the Waikas is Virola
theiodora (Spr. ex Benth.) Warb., as established by voucher
specimens which Bo Holmstedt and I collected from two dif-
ferent groups of these Indians in Brazil in 1967 (Schultes and
Holmstedt, 1968). Other species have been indicated by Ettore
Biocca for the same region (Biocca, 1966), but it seems that
botanical material upon which corroboration of these identifi-
cations can be based is no longer available if, indeed, it ever
was collected: the other species that he reports are V. calophyl-
loidea, V. cuspidata, (Benth.) Warb. and V. rufula (Mart. ex
DC.) Warb. Since these species do give positive alkaloid reac-
tions to spot tests in the field, it is possible that they are actually
used as the basis of narcotic preparations.

Subsequent to the discovery of the use of Viro/a in preparing

315

a snuff, it was learned that the Witoto Indians of Colombia and
their neighbours, the Boras and Muinanes, employ the red
exudate of several species of Virola to make pellets which are
ingested to induce intoxication (Schultes, 1969). In 1969, the
use of V. theiodora for this purpose was first reported from the
Witotos of the Rio Karaparana of Colombia, but these Indians
indicated that several other species might also be so utilized
(Schultes and Swain, 1976). In 1976, with Holmstedt and
Plowman, I learned that the Witotos and Boras of adjacent
parts of the Peruvian Amazon do indeed employ several
species for this purpose. V. elongata, V. loretensis A.C.Sm., V.
Pavonis (DC.) A.C. Sm. and V. surinamensis (Rol.) Warb.
(Schultes, Swain and Plowman, 1977). Of extreme interest was
the report that the Boras apparently employ the myristicaceous
Iryvanthera macrophylla (Benth.) Warb. as the basis of one of
these hallucinogenic preparations.

There is one group of very primitive, nomadic Maku Indians
in the Rio Piraparana of Colombia, who drink the resin of
Virola elongata directly with no preparation in order to induce
the intoxication. This observation, based on voucher speci-
mens, was made by the British anthropologist Peter Silver-
wood-Cope in 1969. There are, likewise, vague suggestions
written on herbarium material collected by the American
botanist Julian Steyermark in 1964, that witch doctors in Ven-
ezuela smoke the inner, dried bark of V. sebifera Aubl. for
curing fevers or to drive away evil spirits. Both V. elongata and
V. sebifera are alkaloidal, and these reports, consequently,
may be based in actual effects of the plants as used.

Species indicated in the foregoing discussion:

Irvanthera macrophylla (Spruce) Warburg in Nov. Act. Nat. Cur. 68 (1897)
15S.

Virola calophylla Warburg in Nov. Act. Nat. Cur. 68 (1897) 231.

Virola calophylloidea Markgraf in Fedde Repert. 19 (1923) 24.

Virola cuspidata (Benth.) Warburg in Nov. Act. Nat. Cur. 68 (1897) 176.

Virola elongata (Benth.) Warburg in Nov. Act. Nat. Cur. 68 (1897) 178.

Virola loretensis A.C. Smith in Bull. Torrey Bot. Club 60 (1933) 95.

Virola Pavonis (A.DC.) A.C. Smith in Brittonia 2 (1937) 504.

Virola rufula (Mart. ex DC.) Warburg in Nov. Act. Nat. Cur. 68 (1897) 181.

Virola theiodora (Spr. ex Benth.) Warburg in Nov. Act. Nat. Cur. 68 (1897)
187.

Virola surinamensis (Rol.) Warburg in Nov. Act. Nat. Cur. 68 (1897) 208.

316

LEGUMINOSAE

Anadenanthera Spegazzini

Historically, the most important South American hallucino-
genic snuff is yopo orniopo, now used almost exclusively in the
Orinoco basin of Colombia and Venezuela.

The earliest report of what undoubtedly must have been this
snuff dates from observations made in 1496 (Safford, 1916).
The Taino Indians of Hispaniola called the snuff cohoba. They
inhaled it to communicate with the spirit world. It was, accord-
ing to anearly writer, **. . . so strong that those who take it lose
consciousness; when the stupefying action begins to wane, the
arms and legs become loose and the head droops .. . and
almost immediately they believe that they see the room turn
upside-down and men walking with their heads downwards.”
Because of the almost complete disappearance of Indians from
the Caribbean, the use of the cohoba snuff no longer exists in
the Antilles. For many years, cohoba was thought to have been
tobacco-snuff, in spite of the clearly very different effects
noted in this earliest observation.

In 1916, the American ethnobotanist William E. Safford
definitely identified the cohoba of the West Indies as the same
intoxicating snuff as yopo of the Orinoco (Safford, 1916). Point-
ing out that the intoxication caused by yopo of the Orinoco was
very similar to that described for cohoba, Safford suggested
that the two must be one snuff under different names. He
reasoned that, since the West Indian islands were populated by
several invasions of Indians from northern South America, the
custom of using this snuff — as well as the source plant itself —
had been introduced to the Caribbean Islands from the Orinoco
basin. The pattern of distribution of this tree in the West Indies
does indeed suggest that it is an introduction.

The snuff was mentioned by the early Jesuit explore of the
Orinoco region, José Gumilla, in his book E/ Orinoco Ilustrado
of 1741.

The earliest botanical report of yopo appears to be that of the
German naturalist-explorer Baron Alexander von Humboldt
who, in 1801, identified the source as Acacia Niopo — an early
name for the plant now known as Anadenanthera peregrina

oy

(L.) Speg. (Humboldt, 1819). In 1819, he reported that the May-
pure Indians of the Orinoco, whom he had seen prepare the
drug eighteen years earlier, softened and kneaded the beans of
this leguminous tree into small cakes with cassava flour (Mani-
hot esculenta Cranz.) and lime from snails; these cakes were
the supply of snuff when it was desired. Von Humboldt erron-

eously reported that “*. . . itis not to be believed that the Niopo
acacia pods are the chief cause of the stimulating effects of the
snuff’? but that ‘*. . . these effects are due to the freshly cal-

cined lime.”’ The botanist K. S. Kunth, who worked with
Humboldt, reported in 1825 the following concerning this
plant: ““Ex seminibus tritis calci vivae admixtis fit tabacum
nobilisque Indi otomacos et guajibos utuntur’’ (Kunth, 1825).

It was the British plant-explorer Richard Spruce, however,
who offered the earliest detailed report on the narcotic, basing
it on field studies amongst the Guahibos of the Orinoco of
Colombia and Venezuela in 1851 (Spruce, 1908). He collected
not only voucher herbarium specimens of Anadenanthera
peregrina but material of the pods and beans for chemical study
and ample ethnological material connected with the prepara-
tion and use of yopo — material still preserved at the Royal
Botanic Gardens, Kew. It is interesting to note that the seeds
collected in 1851 for chemical study were not analyzed for
more than a century and that their analysis was published only
in 1977 (Schultes, Holmstedt, Lindgren and Rivier, 1977).

It is believed that another species of Anadenanthera — A.
colubrina (Vell.) Brenan — was formerly employed in prepar-
ing an intoxicating snuff Known as vilca or huilca in southern
Peru and Bolivia and cébi/ in northern Argentina. This species
of Anadenanthera has the same active principles — tryp-
tamines — as the closely related A. peregrina of the Orinoco
and could have been the source of an inebriating preparation.
The evidence, however, is wholly circumstantial and rather
weak. No voucher material has ever been connected with the
preparation of an hallucinogenic snuff in southern South Amer-
ica, and, of course the Indians who formerly prepared vilca and
cébil have long since disappeared (Altschul, 1967). Recent
studies have, however. discovered the hallucinogenic smoking

318

of A. colubrina amongst Indians of the northwest of Argentina
(Califano, 1976).

At the present time, the use of Anadenanthera peregrina in
elaborating yopo is restricted apparently to the Orinoco Valley
and adjacent regions. There are snuffs prepared and employed
in other areas of tropical South America which have been
attributed toA. peregrina, although, due to the limited distribu-
tion of this species and lack of voucher specimens, there are
serious doubts that these snuffs are, in reality, derived from
this legume. It may possibly be that a century ago, the distribu-
tion of A. peregrina, as aresult of Indian cultivation of the tree,
could have been somewhat wider.

One example of the confusion resulting from incomplete
botanical identification of drugs — or lack of it — are the maps
prepared for anthropological use, in which great areas of South
America are blacked in as regions where snuff from Anadenan-
thera peregrina is employed — even in vast expanses where it
is known that the plant does not occur (Cooper, 1949; Lathrap,
1975). There are a number of other psychoactive snuffs used in
South America — especially tobacco, rapé dos indios, coca,
epena and koribo — yet, because of its greater historical im-
portance, yopo has immediately come to the fore in an-
thropological literature as the major narcotic snuff.

Species indicated in the foregoing discussion:

Anadenanthera colubrina (Vell.) Brenan in Kew Bull. 1955, 2 (1955) 182.

Anadenanthera colubrina (Vell.) Brenan var Cebil (Griseb.) Altschul in
Contrib. Gray Herb. Harvard Univ. 193 (1964) 53.

Anadenanthera peregrina (L.) Spegazzini in Physis 6 (1923) 313.

Anadenanthera peregrina (L.) Spegazzini var. falcata (Benth.) Altschul in
Contrib. Gray Herb. Harvard Univ. 193 (1964) 50.

Mimosa Linnaeus

Here we still have real problems in precise identification,
and it may be too late to solve them.

Although the use of vinho de jurema, an intoxicating drink
once important in aboriginal cultures in northeastern Brazil,
has apparently become extinct, this narcotic preparation was

319

formerly a major hallucinogen. It was the basis, for example, of
the ajuca ceremony of the Pankaruru Indians and was em-
ployed also amongst the Kariri, Tusha, Fulnio, Guegue,
Acroa, Pimentiera, Atanaye and other tribes now totally dis-
appeared or fully acculturated. Furthermore, the magico-reli-
gious use of this Indian narcotic apparently early entered some
of the Afro-American rituals in eastern Brazil.

An early — and probably reliable — report by the Brazilian
writer Caminhoa, quoted by the Brazilian botanist Mello
Morais in 1881, referred to the hallucinogenic effects of jurema.
He stated that medicine men experienced ‘‘fantastic and
agreeable dreams’’ following its ingestion (Mello Morais,
1881).

Another early report by the Brazilian missionary José Mon-
teiro de Noronha suggested in 1768 that the use of jurema may
once have been much wider (Noronha, 1768). Another early
Brazilian report maintained in 1843 that, among the Amanajoz
Indians of the Rio Negro in western Amazonia, warriors,
strong young men and old women singers took a narcotic drink
from the root of a certain tree called jurema. Still another
Brazilian missionary — the Jesuit Joao Daniel — stated that a
number of Amazonian tribes used jurema. These three reports
are most certainly erroneous and confused, since they all were
probably second-hand, and one of them obviously confused
jurema with yopo-snuff and coca powder, both of which are
used only in the Orinoco and western Amazon regions.

Jurema does not occur in the rain-forested Amazon. The
plant or plants yielding vinho de jurema are native only in the
xerophytic caatinga vegetation of the States of Pernambuco
and Paraiba. While it is true — as has been pointed out — that
certain tribes (e.g., the Amanajo) migrated to various points in
the Amazon from northeastern Brazil in the late 1700's, the
major question remains unanswered: where could these In-
dians have found jurema roots in their new and very different
environment? In a weak defense of the early and probably
erroneous reports, we might mention the slight possibility that
the term jurema referred to a number of different psychoactive
plants and that these early migrant Indians applied the term to

320

plants in their new home with properties similar to those of the
jurema that they had known.

Jurema means ‘‘intoxicating drink’? in Tupi-Guarani and,
therefore, was applied probably in a wide sense by the peoples
who used the term.

It was only in 1873 that botanical studies on jurema began
with the work of the Brazilian botanist Aruda Camara who
reported several plants as jurema. A bush known as jurema
branca, which he identified as ‘‘Mimosa Jurema alba’’ was
‘used as a narcotic’. Jurema préta, which he referred to
Acacia Jurema Mart., grew, he stated, only in the driest areas
of the caatingas and sertoes, describing it as a large plant from
which ‘‘the natives prepare a drink which brings on an en-
chantment, transporting them heaven’’.

Later, in 1881, Mello Morais wrote more extensively on the
botanical aspects of jurema. He reported that the “‘country folk
cure fatigue and with the bark of this tree’’ and that “the
Indians extract from jurema a certain kind of intoxicating wine
with delightful effects; to make it, they strip off the bark and,
after boiling it for 24 hours, they add honey to counteract the
astringency of the inebriating drink which is kept for later use.”
Mello Morais identified the source as a new species of Mimosa
which he described and called M. Jurema. It is obvious then
that his observations were based on botanical specimens.

In 1946, the Brazilian biochemist Oswaldo Gongalves de
Lima collected material of the several juremas in northeastern
Brazil. The material was identified by Adolpho Ducke. Jurema
preta, he found, was Mimosa nigra Huber and M. hostilis
(Mart.) Benth. It was the roots of the latter — M. hostilis — that
yielded the *‘miraculous drink’? consumed ritually in the ajuca
ceremony. Jurema branca, used as a stupefacient, 1s M. verru-
cosa Benth. and other closely allied species.

From this authentically identified material of Mimosa hos-
tilis, Gongalves de Lima isolated an alkaloid which he called
nigerine (Goncalves, 1946). The compound was later shown to
be identical with N,N-dimethyltryptamine.

With the knowledge that the active principal of jurema is
N,N-dimethyltryptamine, there arises a further and still uni-

321

vestigated problem. This chemical constituent is believed not
to be active when taken orally, unless it is accompanied by a
monoamine oxidase inhibitor. No such inhibitor has been re-
ported from the roots of Mimosa hostilis. Is there one present
in minute amounts which has not yet been detected? Or per-
chance did the Indians employ as additives parts of other plants
which contain a monoamine oxidase inhibitor? There are here
indicated several courses of investigation: 1) chemical exam-
ination with modern sophisticated techniques of the roots of M.
hostilis, an abundant shrub now of relatively easy access; and,
2) if it still be possible, ethnobotanical field work in northeast-
ern Brazil in the hope that there are still living in the hinterlands
a few elderly Indians knowledgable about the preparation of
vinho de jurema.

Species indicated in the foregoing discussion:

Mimosa hostilis (Mart.) Bentham in Trans. Linn. Soc. 30 (1875) 415.
Mimosa verrucosa Bentham in Hooker Journ. Bot. 4 (1842) 390.

MAPIGHIACEAE

Banisteriopsis C.B. Robinson et Small

Undoubtedly the most widely used and probably culturally
the most important South American hallucinogen is the drink
variously known as ayahuasca, caapi, natema, pindé or yaje.
Employed in the western half of the Amazon Valley in Bolivia,
Brazil, Colombia, Ecuador and Peru, in the uppermost reaches
of the Orinoco in Colombia and Venezuela and even, in spora-
dic localities, on the Pacific slope of the Colombian and Ecua-
dorian Andes, the drug understandably varies significantly in
its preparation from locality to locality and even from tribe to
tribe (Naranjo, 1970).

Made basically from the bark of lianas of the Malpighiaceae,
the major ingredient is either Banisteriopsis Caapi or B. ineb-
rians, the bark of which contains three 8-carboline alkaloids.
Other malpighiaceous plants in this and other genera have,
however, on occasion been indicated as sources of the hallu-
cinogenic drink: B. Cabrerana Cuatr., B. Martiniana (Juss.)

322

Cuatr. var. /aevis Cuatr., B. quitensis, Ndz. Mascagnia glan-
dulifera Cuatr., Tetrapteris methystica R. E. Schult., 7. mu-
cronata Cav. (Schultes, 1957: Friedberg, 1965; Schultes, 1975;
Cuatrecasas, 1958).

The study of the botany and chemistry of this narcotic have
been complicated by the discovery that many other plants in
sundry families are frequently employed as additives, primar-
ily to alter the effects of the drug. Botanical investigations have
been made more difficult by the lack of taxonomic understand-
ing of the Malpighiaceae until recently and chemical analyses
have suffered from both this lack of botanical precision and
from the usual absence of voucher specimens on which exact
identification of the vegetal material might be authenticated.

The earliest report of ayahuasca appears to be that of the
Ecuadorian geographer Manuel Villavicencio who, in 1858
stated, without citing botanical material, that the source of the
drug, was a vine employed by the Zaparo, Angatero, Mazan
and other tribes of the Rio Napo (Villavicencio, 1858).

Seven years earlier, however, in 1851, Spruce had dis-
covered Tukanoan Indians along the Rio Uaupes of Amazo-
nian Brazil using a liana known as caapi, but his observations
were not published until later (Spruce, 1874, 1908). He pre-
cisely identified caapi as a new malpighiaceous species, which
he named Banisteria Caapi. Recent botanical studies have
shown that the concept is more correctly accommodated in the
related genus Banisteriopsis. Spruce made an ample collection
of the liana in full flower — specimens taken from the same vine
from which the drink was prepared. He was so far ahead of his
time, actually, that he collected stems for chemical analysis —
material that was not chemically studied until 1969 (Schultes,
Holmstedt and Lindgren, 1969).

Later, in 1853, Spruce met with the use of caapi amongst the
Guahibos of the Orinoco where, he reported, the Indians “‘not
only drink an infusion, like those of the Uaupes, but also chew
the dried stem”’ (Spruce, 1908). Again, in 1857, he encountered
the Zaparos of Ecuador taking ayahuasca, and he considered it
to be ‘‘the identical species of the Uaupés, but under a different
name’ (Spruce, 1908).

In the century that followed Spruce’s remarkable work,

323

many explorers, botanists and others — von Martius, Orton,
Crevaux, Koch-Grunberg, to name only a few — referred to
ayahuasca, caapi, or yaje, usually quite casually and without
citing botanical material: stating merely that the drug came
from ‘‘a jungle liana’’ (Schultes, 1957; Friedberg, 1965).

Of outstanding interest was the work in 1922 of Rusby in
Bolivia. Rusby vouchered all of his collections of drug plants,
but we have been unable to locate a specimen of Banisteriopsis
in his material (Rusby, 1923). Another contribution of major
importance was the publication in 1931 by the American
botanist Conrad V. Morton of the detailed field notes made by
the famous Peruvian botanical collector Guilielmo Klug in the
Colombian Putumayo (Morton, 1931). The second of the two
major sources of the hallucinogenic drink, Banisteriopsis ineb-
rians, was based on Klug’s collection. Similarly, the collec-
tions of the Russian botanists Varanof and Juzepezuk in the
Caqueta area of Colombia in 1925-26 added valuable data based
upon botanical material (Schultes, 1957). The field work of the
Colombian botanist Hernando Garcia-Barriga has added ap-
preciably to our knowledge (Garcia-Barriga, 1958). In 1957, I
published a review of the then known sources of the hallucino-
gen, incorporating my own field studies in the Colombian
Amazon witha review of the literature that had grown up onthe
problem (Schultes, 1957). This was followed in 1965 by an
exceptionally complete ethnobotanical survey of the drug by
the French ethnobotanist Claudine Friedberg, who had carried
out field work in Peru (Friedberg, 1965). In the meantime, the
taxonomist José Cuatrecasas of the Smithsonian Institution
had provided, for the first time, a firm basis for classification of
ethnobotanical and phytochemical problems with his mono-
graph of the Malpighiaceae of Colombia, published in 1958
(Cuatrecasas, 1958).

But serious complications early arose in attempts to identify
the hallucinogen. In 1890, a missionary confused the tree-
species of Datura or Brugmansia employed by the Jivaro with
their malpighiaceous narcotic — a confusion that quickly
entered the pharmacological and chemical literature. Unfortu-
nately, it has persisted. And at one time, yaje was attributed
even to the Aristolochiaceae (Rouhier, 1924)!

324

Then there arose, primarily from uncritical interpretation of
Spruce’s field notes, an extraordinary presumption that gained
very wide acceptance in the literature: it presumed that, al-
though ayahuasca and caapi were derived from Banisteriopsis,
yajé referred to the apocynaceous Prestonia amazonica
(Benth.) Macbr. Although this suggestion has been thoroughly
discredited (Schultes and Raffauf, 1960), it persists in the popu-
lar and, unfortunately, even in the technical literature. It is
responsible for much of the confusion in chemical and phar-
macological studies. Spruce, in his field notes, suggested that
the Indians of the Vaupés sometimes added a plant, which he
felt might have been what the Indians called caapi-pinima
(‘painted caapi’’): he noted that it looked like the apocynace-
ous plant that he had collected earlier in the central Amazon
and which was described as a new species: Haemadictyon
amazonicum Benth. ex Muell.-Arg., now known as Prestonia
amazonica (Spruce, 1908). This tentative observation was later
uncritically interpreted by a French researcher who had never
been in the field as a flat assertion and was so published
(Reinberg, 1921). Prestonia amazonica, collected but once in
over 125 years, is known only froma locality nearly 1500 miles
from the Uaupés; with all of the now known admixtures used
with the drink, there is no question that the local additive to
which Spruce referred could not have been Prestonia amazon-
ica, especially since so many botanists and ethnologists have
worked in the region where Spruce collected and have not
found a species of Prestonia employed for this hallucinogenic
purpose.

It is now definitely known that the major constituents of
ayahuasca, caapi and yajé are species of Banisteriopsis. There
remains, however, the colossal task of elucidating the many
additives which Amazonian Indians put with the hallucinogen
(Schultes, 1972). These additives or admixtures may be very
localized — even to a single medicine man — or they may be
widely utilized. There is, in many instances, good reason to
assume that they drastically alter or strengthen the effects of B.
Caapi or B. inebrians alone.

The Siona of Colombia add what is probably Brugmansia
suaveolens (H. et B. ex Willd.) Bercht. et Presl., in itself an

325

hallucinogen. Their neighbors, the Inganos, add the amaran-
thaceous Alternanthera Lehmannii. The Kofan and Jivaro
tribes value the solanaceous Brunfelsia as an admixture.
Makuna medicine men in the Colombian Vaupes add a few
crushed leaves of the apocynaceous Malouetia Tamaquarina
A.DC. Tobacco is occasionally used as an admixture
(Schultes, 1957). Several lianas, without identification, are said
to be added, but it has not been possible to identify them for
lack of specimens. A very thorough field and laboratory study
of plant additives to ayahuasca by the Swiss biochemist Lau-
rent Rivier and the Swedish chemist Jan-Erik Lindgren has
disclosed the use of at least two dozen plants, including ferns
and several cactuses (Rivier and Lindgren, 1972).

Recently, two of the many admixtures have attracted special
attention, because phytochemical investigation of botanically
vouchered material has substantiated folk uses. More often
than not, specialists have explained away indigenous uses of
plants as being grounded in superstition. These two admixtures
in question are Banisteriopsis Rusbyvana (Ndz.) Morton and
Psychotria viridis R. et P. The leaves — not the bark — of both
plants are used, apparently never together, however, as admix-
tures.

In 1965, the French chemist M. J. Poisson reported that the
leaves of Banisteriopsis Rusbyana contained relatively high
concentrations of N,N-dimethyltryptamine (Poisson, 1965).
This discovery, corroborated by several later investigators
(Agurell, Holmstedt and Lindgren, 1968; der Marderosian et
al., 1968) was the first report of a tryptamine in the Malpighia-
ceae. There were also minor tryptamines present, as well as
traces of B-carbolines. The leaves of B. Rusbyana, known in
Colombia and Ecuador as oco-yajé, are said to heighten and
lengthen the visual hallucinations of the intoxication. It is now
clear that there is a chemical basis for this use: the resulting
drink, containing the 6-carboline alkaloids of B. Caapi and B.
Rusbyana — both hallucinogenic indole derivitives — 1s, in
effect, far more potent.

More recent field work has indicated the use as additives in
widely separated parts of the Amazon of the leaves of several
species of Psychotria — especially P. viridis (der Marderosian,

326

1970). This additive, which is apparently far more commonly
utilized than Banisteriopsis Rusbyana, likewise has been found
to contain N,N-dimethyltryptamine — the first indication of
this typtamine in the Rubiaceae.

Several writers — notably Spruce and Koch-Grunberg —
mentioned more than one kind of caapi in the basin of the Rio
Uaupés of Colombia and Brazil. In 1948, I witnessed the prepa-
ration and experimented with a caapi prepared from a different
malpighiaceous plant amongst the nomadic Maku Indians of
the Rio Tikié, in northwesternmost Brazil. Specimens of the
liana, from the bark of which a cold water infusion was made,
represented an undescribed species of the genus Tetrapteris,
closely allied to Banisteriopsis, which I described and named
T. methystica (Schultes, 1954c). It is worth considering the
possibility that this species is the second ‘kind’ of caapi
mentioned by Spruce and Koch-Grtinberg, and it might be that
the name caapi-pinima (‘‘painted caapi’’) reported by both of
these investigators alludes not to coloured leaves but to the
unusual yellowish hue of the drink in contrast to the normal
dark brown colour of the hallucinogen prepared from the
species of Banisteriopsis.

Much field and laboratory research must be carried out
before a complete understanding of the ayahuasca-caapi-yaje
complex is available. It is disconcerting that the rapidity of
acculturation throughout the Amazon Valley may soon bring
about the loss of much of the knowledge of plant uses amongst
the Indians. It will then be too late to retrieve extremely valu-
able information.

Species indicated in the foregoing discussion:

Bonisteriopsis Caapi (Spr. ex. Griseb.) Morton in Journ. Wash. Acad. Sci. 21
(1931) 485.

Banisteriopsis Cabrerana Cuatrecasas in Webbia 23, pt. 1 (1958) 493.

Banisteriopsis inebrians Morton in Journ, Wash. Acad. Sci. 21 (1931) 485.

Banisteriopsis Martiniana (Juss.) Cuatrecasas var. laevis Cuatrecasas in
Webbia 13 (1958) 502.

Banisteriopsis quitensis Niedenzu in Lect. Lyc. Brunsberg (1900-01): 10
(1900) Malpigh. 427.

Banisteriopsis Rusbyana (Ndz.) Morton in Journ. Wash. Acad. Sci. 21 (1931)
487.

Muascagnia glandulifera Cuatrecasas in Webbia 13 (1958) 365.

a |

Tetrapteris methystica R.E. Schultes in Bot. Mus. Leafl., Harvard Univ. 16
(1954) 202.
Tetrapteris mucronata Cavanilles, Diss. 9 (1790) 434, t. 262.

SOLANACEAE
Latua Philippi

There is an unusual and rare spiny shrub or small tree —
apparently a strict endemic of the coastal mountains of central
Chile — which has long been used as a virulently toxic plant
capable of producing delirium, hallucinations and, occasion-
ally, permanent insanity. The genus is monotypic, consisting of
a single species: Latua pubiflora (Griseb.) Baillon.

Although the identification of this hallucinogen has been
available for more thana century, little, until recently, has been
known of its use and properties.

Latua pubiflora was first described by the German botanist
August H.R. Griesbach in 1854 as a species of Lycioplesium.
Later, in 1858, the Chilean botanist R. A. Philippi described the
plant as representing a new genus, Latua, giving it the name
Latua venenosa (Philippi, 1858). It was not until 1888 that the
French taxonomist Henri E. Baillon made the correct combi-
nation, assigning the currently accepted name of Latua pubi-
flora.

The narcotic use of Latua pubiflora — known locally as latue
or arbol de los brujos (*‘sorcerers’ tree’’) — was first associ-
ated with the plant in 1858 by Philippi: **. . . the native Indians
posses a secret to madden a person by means of a vegetable
poison’’. He related several cases of intentional and accidental
intoxication (Philippi, 1858).

A few collections of Latua pubiflora have been made during
the past 120 years, but a thorough survey of the plant, and its
use, following extensive field work, has only recently been
published (Plowman, Gyllenhaal and Lindgren, 1971). The oc-
currence and use of latue is still a closely guarded secret sur-
rounded by superstition, as it is utilized in magico-medical rites
by shamans and sorcerers.

Species indicated in the foregoing discussion:
Latua pubiflora (Griseb.) Baillon, Hist. Plant. 9 (1888) 334.
328

CONCLUSION

A few closing words may bring into relief several salient
points of interest.

1) There is no doubt — whether or not supporting archaeol-
ogical or early literature evidence be available — that the
indigenous use of all of the major psychoactive drugs of South
America indicates great age.

2) For coca, archaeological material puts its use back at
least as far as 3000 BC — 5000 years ago. Actual authenticated
botanical identification came about in 1749 — 229 years ago.

3) The earliest literature reference to yopo-snuff is appar-
ently 482 years old — made in 1496. The source plant was not
botanically determined until 1801.

4) For the hallucinogenic preparations made from Virola,
the first literature reference seems to be one published in 1923.
The actual source plants, however, were not identified specif-
ically on the basis of botanical material until 1954.

5) A description of the use and effects of ayahuasca ap-
peared in the literature in 1858, although the same drug, under
the name caapi, had been botanically collected, named and
described, but not published, in 1851.

6) The Chilean arbol de los brujos or latué was discovered
and botanically identified in 1854, but the narcotic use of the
species was indicated only in 1858.

7) The earliest literature reference to jurema dates from the
late 1700’s. Botanical identification began in 1873 and was
substantiated only in 1946.

These observations indicate how much basic botanical work
has been accomplished during the past century. This century
has been marked by ever increasing acculturation and even
disappearance of native cultures in South America. There are
sundry minor yet extremely interesting hallucinogens and nar-
cotics — half a dozen of them reported only during the last 10 or
15 years — of which we know next to nothing. It behooves
ethnobotanists and enthnopharmacological investigators to
study these psychoactive plants, their use and their signifi-
cance in aboriginal societies before the race with acculturation
or extinction of indigenous cultures is forever lost.

Azo

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Friedberg, C. 1965. Des Banisteriopsis utilisés comme drogue en Amerique
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780.

Garcia-Barriga, H. 1958. El yajé, caapi o ayahuasca: un alucinégeno
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Gongalves de Lima, O. 1946. Observagoes sobre 0 vinho da jurema utilizado
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Gumilla, J. 1741. El Orinoco Mustrado. Manuel Fernandez, Madrid. Pp.
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von Humboldt, A. 1819. Voyage aux Régions Equinoxiales du Nouveau
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Koch-Grunberg, T. 1923. Vom Roroima zum Orinoco. Verlag Strecker und
Schroder, Stuttgart. 3: p. 386.

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G. Leverault, Paris. p. 20.

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300 B.C. Field Museum of Natural History, Chicago.

der Marderosian, A., K. M. Kensinger, J.-M. Chao and F. J. Goldstein. The

330

use of hallucinatory principles of a psychoactive beverage of the
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L. Garnier, Rio de Janeiro. p. 228.

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ao)

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Tropicale commentationes XII]a. Further notes on Virola as an orally
administered hallucinogen. Journ. Psyched. Drugs 8: pp. 317-324.

Schultes, R. E., T. Swain and T. Plowman. 1977. De plantis toxicariis e
Mundo Novo tropicale commentationes XVII. Virola as an oral hal-
lucinogen among the Boras of Peru. Bot. Mus. Leafl., Harvard Univ.
25: pp. 259-272.

Spruce, R. 1874. On some remarkable narcotics of the Amazon Valley and
Orinoco. Geogr. Mag., n.s. 1: 184-193.

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Villavicencio, M. Geografia de la Republica del Ecuador. R. Graighead,
New York.

332

PLATE 36

620 LIVRE VIII.

mode de nutrition. Nous n’apprendrons jamais a digércr des terres et 4 les
assimiler ; mais, depuis que les grands travaux de MM. Gay-Lussac et Thénard
nous ont fait connoitre que de légéres différences de proportions d’oxigéne,
d’hydrogéne et de carbone caractérisent seules le bois le plus dur et la matiére
amylacée , comment nier que la chimie ne puisse réussir un jour a convertir en
substance alimentaire ces énormes masses végétales, ces tissus a fibres endurcies
qui composent le tronc des arbres de nos foréts? Pour qu'une telle découverte
fit importante, elle devroit étre fondée sur des procédés simples et peu couteux :
mais dans cette supposition, qui ne paroit guére probable, elle changeroit
organisation des corps politiques, le prix du travail, la-distribution de la
population sur le globe. En rendant homme plus indépendant, elle tendroit a
dissoudre les liens de la société , a sapper les bases de l'industrie et de la civilisation.

Le petit village d'Uruana est plus difficile & gouverner que la plupart des
autres missions. Les Otomaques sont un peuple inquiet, bruyant, effréné dans
ses passions. Ils n’aiment pas seulement avec exces les liqueurs fermentées de
manioc et de mais et le vin de palmier; ils se mettent aussi dans un état parti-
culier d'ivresse, on pourroit presque dire de démence, par l’'usage de la poudre
de nzopo '. Ils cueillent les longues gonsses d'une Mimosacée que nous avons
fait connoitre sous le nom d’ Acacia Niopo ”; ils les mettent en morceaux, les
humectent et les font fermenter. Lorsque les graines amollies commencenta noircir,
ils les pétrissent comme une pate ; et, aprés y avoir mélé de la farine de manioc et
de la chaux tirée de la coquille d'une Ampullaire, ils exposent toutela masse a un
feu trés-vif'sur un gril de bois dur. La pate durcie prend la forme de petits gateaux.
Lorsqu’on veut s’en servir, on la réduit en une poudre fine qu’on place sur un plat
ce 5 ou6 pouces de largeur. L’Otomaque tient ce plat, qui a un manche, dans sa
roain droite, tandis qu'il aspire le niopo par le nez a travers un os fourchu
doiseau dont les deux extrémités aboutissent aux narines. L’os, sans lequel l'Oto-
maque ne croiroit pas pouvoir prendre cette espece de tabac en poudre, a
7 pouces de longueur : il m’a paru ¢tre le tarse d’un grand Echassier. J'ai
envoyé le niopo et tout ce singulier appareil a M. de Fourcroy a Paris, Le
ntopo est si excitant que les plus petites portions font éternuer violemment ceux

' En maypure, nupa ; les missionnaires disent Aopo.

* Cest un Acacia a feuilles trésdélicates, et non un Inga, comme a dit par mégarde M. 7¥ilidenow ( Spec.
Plant., Tom. 1V, Pl. 11, p- 1027). Une autre esptce de Mimosacée que nous ayons rapportée (le Chiga des
Otomaques et le Sip des Maypures ) donne des graines dont la farine est mangéc 4 Uruana comme du manioc.
(est de cette favine que Von prépare le pain de chiga qui est commun & Cunaviche et sur les bords du Bas-

Orcnoque. Le Chiga est une espice d’ nga, et je ne connois point d’autre Mimosacée qui supplée aux Céréales.

Plate 36. Von Humboldt’s identification of the source of the narcotic snuff
niopo of the Otomac Indians.

333

PLATE 37

“
16. Jahrgang. m2 32. 13. August 1858.

BOTANISCHE ZEITUNG.

Redaction: Hugo ron Mohl. — D. F. L. con Schlechtendal.

Inhalt. Orig.: Philippi, Latua, cin neues Genus der Solanaceen u. einize Reisebemerkungen. Lit. :
Verhandl. d. Vereins f, Naturkunde z. Presburg. Hl. — Nordlinger, 50 Quers hnitte d. in Deutschland
wachsend, hauptsachl. Hélzer. — Schwiztein, Analysen z. d. natirl, Ordn., der Gewachse in Europa, —
Max Schultze, Innere Bewegungserscheinungen b. Diatomeen d. Nordsee. — Kotsehy, Reise in den
eilicisehen Taurus. — Hannaford, Jottings im Australia, — K. Not.: Scheeffer, tb. Baumwollenfaser

Latua Pb.. ein neues Genus der Solanaceen. dessen Rinde Ausserlich und inuerlich in Gestalt ei-
nes Dekoktes fiir ein vortreffliches Mittel yegen
Quetachunyen, Erschitterungen durch Fall, Huf-
schlage u. s. w. gilt. So theilte er mir unter an-

Von
Prof. Dr. HB. A. Philippi in santjago.

Bereits vor sechs Jahren erfuhr ich in der Pro- | dern folgenden Fall mit, der kurzlich passirt war
ving Valdivia, dass die dortigen Indiaver das Ge- | Finer seiner Holzarbeiter hatte einen heftigen Schlag
heimniss besissen, vermittelst eines vegetabilischen | mit dem stumpfen Ende einer Axt erhalten, und ging
Giftes die Menschen verriickt zu machen, und zwar in den Wald, um sich dagegen Hinde von Tayu zu
auf ldngere oder kiirzere Geit, je mach der Starke holen, ergriff aber statt dessen Latue und trank
der Gabe dieses Giftes, dass sie die Sache aber als | die Abkochung von diesem Gift. Fast uumittelbar
ein grosses Geheimniss behandelten. Dem Pater | darauf wurde er verriickt und lief in die Walder,
Romualdo, Missiondr in Daglipulli, gelang es zu er- | wo er erst nach Ablauf von drei Tagen in bewusst-
fahren, dass die Pflanze ein hoher Strauch sei, La- losem Zustande gefunden wurde. Kinige Tage ge-
tue heisse, hie und da im Urwalde des Kustengebir- nigten zwar, ihn wieder herzustellen, allein er be-
yes wachse, und endlich auch cinen Zweig zu er- hielt Monate lang heftige Kopfachmerzev. Eben 80

halten. Dieser war jedoch ohne Blatter, indem der ziftig wie die Rinde sind die Friichte, welche durch
Indier, welcher ihn brachte, unstreitig glaubte, der | ihre Farbe und Grosse einige Aehnlichkeit mit jun-
Pater wollte denselben haben, um seine giftigen Ei- | gen Aepfeln haben. Kin Paar Leute, welche auf
dem Weze von Osorno nach Moullin, wo demnach

genschaften, welche hauptsachlich in der Rinde
sitzen, zu ertorschen. Sp&ter erfubv ich von Herrn | die Latue auch wachsen muss, sich verleiten lies-
Jnan Renous Nadheres uber den Latue. Der Strauch | sen, ein Paar Frichte zu essen, kamen fast ohne
sei dem Yuyu oder Palo Santo, Flotowia diacan- | Bewusstsein und yanz verstort an letzterem Orte
thoides, im Wuchse, in dev Dornen und Bl&ttern
iiberaus &hnlich, die Blithe sei aber wie von der Ge-
stalt und Groésse der von Sarmienta repens KR, et P. | vorbringt, und uber die Gegenmittel, welche die In-
(einer Gesneriacee, welche zwischen den Moosen dianer auwenden, um die ubelo Folgeu der Vergil-
und Farrnkrdutern der Stdmme und grésseren Aeste | tung zu beseitigen.

an. Leider habe ich nichts Ndheres ber die Syimp-

tome erfahren, welche der Genuss der Latue her-

kriechend und durch ihre scharlachrothen Blumen Meine Begierde, eine so merkwirdige Pflanze
damit lebhaft contrastirend zu den schénsten Zier- | kennen zu lernen, war natdrlich sehr gross, und

den der Walder des siidlichen Chiles yehdrt), Von | ich sparte keine Miihe, mir Exemplare derselbeun mit
der Frucht wusste mir Herr Renous nichts zu sa- | Blithen und Frucht zu verschaflen, aber lange wa-
gen, wohl aber theilte mir derselbe mehrere FAalle | ren meine Anstrengungen vergehens, Im Jahre 1854
von absichtlichen und unfreiwilligen Vergiftungen | fand Herr C. Ocsenius am Ufer des Riobueno oder
mit. Letztere sind um so leichter, als der Strauch, | Trumao zwischen den Orten los Jantas und Trini-
wie eben gesagt, dem Tayu so sehr dholich ist, | dad einen ihm unbekannten Strauch in Blithe, brach
33

Plate 37. The report by Philippi of the toxic nature of Latua (1858).

334

PLATE. 35

( 20 )

2) Inermes.

30. A. GuACHAPELE. +

A. inermis; foliis bipinnatis; pinnis 4 - 5- jugis ; foliolis
§- &-jugis , obovatis vel subelliptico-oblongis, membra-
naceis, utrinque pubescentibus; glandula in medio petioli
interque summum pinnarum par ; floribus axillaribus, ca-
pitatis , monadelphis; leguminibus linearibus, longe rostra-
tis. (VI. p. 281.)

Guachapele Guayaquilensium.

Crescit in sylvis , prope Guayaquil Quitensium. t Flo-
ret Martio.

Lignum ad construendum aptissimum.

Acacie Lebbeck yalde affinis.

31. A. PECTINATA. |

A. inermis; foliis bipinnatis, eglandulosis; pinnis sub 15-
jugis; foliolis 50-60 -jugis, linearibus, obtusis , suabmucro-
natis , dimidiato -subhastatis, uninerviis, subcoriaceis, supra
glabris , nitidis, subius sericeo-pubescentibus. (WZ. p. 282.)

Inga pectinata. Willd. Sp. pl. 4. p. 1126. «

Cresctt ad ripam flumints Cassiquiares. (Missiones
del Alto Orinoco.) +} Floret Mazo.

An vere hujus generis ?

32. A. Niovo. +

A. inermis ; foliis bipinnatis ; pinnis sub 23 - jugis; fo-
liolis 50- 70- jugis, linearibus, subfalcatis , acutis, obsolete
uninerviis , membranaceis, glabris, ciliatis; glandula supra
basim petioli et inter summum pinnarum par; leguminibus
linearibus, rostratis. (J. p. 282.)

A. Niopo. Humb. Relat. hist. II. p. 620 — 623.

Inga Niopo. Willd. Sp. pl. 4. p. 1027.

Niopo incolarum.

Crescit prope Maypures, Atures, etc. (Mussiones del
Orinoco.) + Fructificat Majo.

Ex seminibus tritis calci vive admixtis fif tabacum nobile quo
Indi oe et Guajibos dicti utuntur. (Humb. Relat. hist. II.
p- 620.

Plate 38. Kunth’s report of the source of the snuff known as niopo. Synop-
sis Plantarum... of 1825.

238

PLATE 39

228 BOTANICA BRASILEIRA

Jurema. (Mimosa jurema.) — Arvore de mediana gran-
deza, que vegeta em terrenos fracos e seccos.

USOS MEDICINAES
A entre-casca da jurema serve em cozimento, para lavar feri-
das. A casca da raiz, serve para curar as diarrhéas, tomado

o cozimento em clyster.
Os sertanejos, curam 0 cansaco, e a cachexia, com a casca

desta arvore.

Os indios extrahem da jurema, certa especie de vinho, que
embriaga, com transporte delicioso; e para este fim, tiram a
casca, poem-n’a em infusio, por 24 horas, coam depois a infusao,
ajuntam-lhe mel de abelhas, para corrigir o gosto adstringente
dessa embriagante bebida, e guardam-n’a para o uso,

Os sertanejos empregam a entre-casca da jurema, como forti-
ficante do apparelho uterino.

Jurubeba. (Solanum paniculatuin.)—Planta, de granudeza
mediana, tendo as folhas largas, e todas ellassemeadas de espiahos.
E’ mui conhecida na Bahia, Sergipe, Alagoas e Pernambuco, onde
vegeta em abunda:cia. Esta planta ¢ mui estimada por suas vir-
tudes medicinaes.

USOS MEDICINAES

A raiz cozida, e dada a beber, cura as gonorrhcas, com-
batendo ao mesmo tempo o virus syphilitico. A fructa, que é
amarga, & grande remedi» para as apvsthemas iaterias; bem
como servem-se do suino bebido, e posto extcruameute nas con-
tusdes, pela manha, e 4 noite, misturado com agua. E’, além
disso, estomacal, e um precioso desobstruente do figado.

O padre-mestre José Bonifacio Bezerra de Mello (meu professor
de latim), constantemente comia, como regalo,a fructa da jurubeba.

Jussara ou assaly. (Alagoas.)—lEspecie de coqueiro,
mui abundante nas Alagoas.

USOS NA INDUSTRIA
A industria péde obter desta planta, optimo vinho.
Jussara.—Especie de capim, cuja folha imita ao capim
Sapé, e deita flor amarella.

Plate 39. The report of jurema made by Mello Morais in 1881 in his Phyto-
graphia ou Botanica Brasiliera.

336

PLATE 40

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vied BISY Ud S9]¥jJWaO SO] USAIIS OS [Bud Bl ep ‘oidg op
uorisoduioo vy oovy onb soy B Sos0l1oyUT UO OU BpIqed BISe
ap sojoajo Soy :soroZnu svy eu souaaol Anu soy v eyutsed
as ou wospnyvhp [yA “AvU0) aqop onb souolBulULd}op SB]
v]So1v souorotjsiodns sns UNDos 1 ‘soUOISIA SB] TIA OPUBNO
oan} onb sopasonoer sop efooor oyens ouI[N [9 Opesed
‘sous yuu sosjoodse soy y oorp onb svyvavsq sey Bins vy
WOS $dI[¥} :SdJUBJSUNIIIO SO] Op OUNDSUIU BpIA UOd viivdeose
ou zaa je} “vue vaommbyend op vaviopody os 1 pe}0q!]
uo vivfop of os 1s onb aod {vovwmey wun ua o}UdUIo}IONJ
oprejolus vied voseo ugyso onb svysnqor seucssed op ¥z1s900U
oaad azooxjd aod saooa svypnut vospnyvhy jo Buoy ofeayes
Tq ‘[eroueS avysa yeu saoaa svunsye 1 BzoqBo ap zopesed
LA0]op od BqRpsOd.I ond Jap, OYoNS op o}LeNj UOLOBSUIS
vie} wpindes uo {erpuajop ou onb sul op ‘seioy svunde
op Opyeuooe t onbsoq un uo opruopunge vquinsy oul odon]
{soust[[aq soyolqo sozjo 1 sanbavd sosoulloy ‘sa110} sEpBAd]d
‘sapepnio sopuvsd ‘sesororep seu svanoodsead sey erqiosed
Opssnder onb Ja ua oa198 oferta un OSany| ‘ezoqvo op soapol
opyues ey vospnynhy jo opvuto} oy opueno onb soep 98
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f 2[qupeide op saqus uvosep sotfe enb oj ueorwnui0o
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IVZUIUIOD | OATB [BV OSIBAITA OP UOLOsUES BT OFoNT ‘ez0quo
dp soopor f sopryea udjzusis fuvjzoidsap os sopeznoRy
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rey0s0 B osuflarp aoaivd uoloov ng “sousWOUOJ sores sBUT
so] tonpoad ¥ wzardura sojyuswo0Ur soood ¥ 1 ‘esiouodns oqep
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SOIPUL SO] SOPO} UOqoq OT S8d9A SETPNUI I seuBld SOT LeTSOIIB
9 sujsondsei sel rep oqop onb orput Ja eqaq Oy 1 OJUaTUIIO0O
soft] un usavy [end Jap (svMTe O OJZoNU op CoNfaq) DoEDnY
vhy opeurry, oonfoq un uvui0, :ayuatnais of ud oysIsU0d
uo1viedo wy ‘saaZnui sus op ow Jap esrBIOIOI0 Baed
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tion of the use of ayahuasca in Ecuador

Villavicencio’s descrip

Plate 40.
(1858)

337

BOTANICAL MUSEUM LEAFLETS VoL. 26, No. 9-10
NOVEMBER-DECEMBER 1978

THE CORRECT NAME FOR SIBERIAN GINSENG

7

DJAJA D. SOEJARTO!,2 AND NORMAN R. FARNSWORTH!

‘*Siberian Ginseng’’ is a vernacular name associated with an
araliaceous plant and its product, which is now imported
primarily from the Soviet Union and marketed in health food
stores here in the U.S.A., Europe and elsewhere. The product
is prepared from the roots of Eleutherococcus senticosus
(Rupr. & Maxim.) Maxim., which is commonly referred to as
‘‘Devil’s Shrub’’, ‘‘Touch-me-not’’, ‘‘Prickly Eleuthero-
coce’’, ‘‘Eleutherococc’’, and ‘‘Siberian Ginseng”. This
species grows in Eastern Siberia, Sakhalin Island, Japan,
South Korea, Manchuria, and the Chinese Provinces of Shansi
and Hopei. A taxonomic description of E. senticosus is pro-
vided by Poyarkova (1973).

It is interesting to note that E. senticosus is not mentioned to
any extent in the early Oriental medical writings as having
useful properties. However, extracts of the roots of E. sentico-
sus have been studied extensively in the Soviet Union for about
25 years: more than 200 scientific periodicals can be found onin
vitro, in vivo, and in situ effects, as well as scores of papers
reporting results of human studies. These studies have been
pioneered by Brekhman (1969, 1970), Dardymov (1976) and
their co-workers, and are continuing. The major effects attrib-
uted to extracts of E. senticosus roots relate to an anti-stress
phenomenon, called by Russian workers ‘‘an adaptogenic ef-
fect’’, the mechanism for which remains unclear.

Since this plant is an important item of commerce, and
because a great deal of confusion seems to be apparent in the
herb industry relative to the priority of the binomial Eleuthero-
coccus senticosus over Acanthopanax senticosus (or vice

' Department of Pharmacognosy and Pharmacology, College of Pharmacy, University
of Illinois at the Medical Center, 833 South Wood St., Chicago, Ill. 60612.

2Research Associate, Field Museum of Natural History (Botany), Chicago; Honorary
Research Fellow in Ethnomedicine, Botanical Museum of Harvard University.

339

versa), we undertook a study to clarify the taxonomic and
nomenclatural position of this taxon.

The genus Eleutherococcus> was established by Maximo-
wicz in 1859 with E. senticosus as the type species, on the basis
of plant specimens collected from Amur and Ussuri (Siberia).
Previously, this plant was known to Ruprecht and Maximo-
wicz as Hedera senticosa Rupr. & Maxim. (1857), but later
Maximowicz recognized it as belonging to a new taxon, for
which he introduced the name Eleutherococcus, and re-named
the plant Eleutherococcus senticosus (Rupr. & Maxim.)
Maxim.

The genus Acanthopanax* was established by Miquel in
1863, when he raised the subgenus Acanthopanax Decne. &
Pl., of the genus Panax L., into a generic status. A new taxon,
Acanthopanax spinosum Miq., accompanied the generic de-
scription.

In 1898, Harms merged Eleutherococcus into the genus
Acanthopanax, and the name of our plant became Acantho-
panax senticosus (Rupr. & Maxim.) Harms. After 1898, some
authors continued to use the binomial FE. senticosus (e.g.,
Nakai, 1924; Liou, 1959: including the references cited there-
in), while others used A. senticosus (e.g., Schneider, 1912;
Harms & Rehder, 1916; Rehder, 1922; Li, 1942).

The union of the genera by Harms demonstrates that a close
taxonomic relationship exists. Subsequently, Harms treated
members of Eleutherococcus under the Section Eleutherococ-
cus of the genus Acanthopanax, whereas members of Acan-
thopanax were treated under the Section Ewacanthopanax.
The creation of these two Sections within the genus clearly
indicates that Harms did recognize the differences that sepa-
rate the two plant groups.

In 1924, Nakai re-instated the generic status of Eleuthero-
coccus, but in 1942, Li treated Eleutherococcus again under
Acanthopanax, as did Harms (1898). In 1950, once again, the
genus Eleutherococcus was re-instated in the Flora of the
U.S.S.R. (Poyarkova, 1973).

After studying specimens belonging to Eleutherococcus and
iPro the Greek pleuthovens = free, and coccon = seed.

4From the Greek akanthos = thorn or spine, and Panax = name of genus.

340

Acanthopanax*, and analyzing the published original descrip-
tions of the same, we came to the conclusion that the major
features that separate the two taxa are as follows:

Eleutherococcus: Flower pedicels not jointed,
flowers in solitary umbels, ovary 5-loculed
and 5-styled (rarely 3-6), fruits globose or
acute with 5 seeds (rarely 3-6).

Acanthopanax: Flower pedicels jointed, flowers in
racemose or paniculate umbels, ovary
2-loculed and 2-styled (rarely 3-loculed and
3-styled), fruits flat or compressed, with 2
(rarely 3) seeds.

Basically, these are criteria used by Bentham & Hooker f.
(1867) in maintaining the separation of Eleutherococcus from
Acanthopanax at the generic level. It is also worthwhile to note
that Nakai (1924) emphasized the differences that separate
these two taxa in the number of styles and the number of cells
(locules) of the ovary, as well as in the shape of the fruits. The
presence or absence of articulation in the flower pedicels,
according to Nakai (1924) is not an important character, since
this phenomenon is seen only in dried specimens, caused by
the unequal shrinking of the parenchyma cells.

We feel that the criteria defined above are strong enough to
separate the two taxa at the generic level. Union of the taxa
results in a heterogeneous group taxonomically, as Li (1942)
has admitted. Li further states that this ‘‘genus [Acantho-
panax, s.l.| is heterogeneous and may ultimately require cer-
tain segregations’’ (1942: 6). On the basis of the present evi-
dence, we recognize the genus Eleutherococcus (with about 15
species) as distinct from Acanthopanax (with about 25
species).

‘We are grateful to the following individuals and institutions for the loan of specimens
of Eleutherococcus and/or Acanthopanax, for examination: Arnold Arboretum, Har-
vard University, Cambridge, Mass.; Field Museum of Natural History, Botany De-
partment, Chicago, IIl.; Dr. Tchang Bok Lee, the Kwanak Arboretum, Seoul National
University, Suwon, South Korea; Professor I.1. Brekhman, Far East Scientific Cen-
tre of the USSR, Academy of Sciences, Vladivostok, U.S.S.R.

34]

As aconsequence, the correct name for **Siberian Ginseng’
should be Eleutherococcus senticosus (Rupr. & Maxim.)
Maxim. The binomial Acanthopanax senticosus (Rupr. &
Maxim.) Harms is thus relegated to synonymy.

The legitimacy of Eleutherococcus senticosus does not
change, even if Eleutherococcus and Acanthopanax are
united. When Harms (1898) merged Eleutherococcus with
Acanthopanax, the name Eleutherococcus should have been
retained, in accordance with the law of priority. Article 57./ of
the International Code of Botanical Nomenclature (Stafleu,
1978) states: “‘When two or more taxa of the same rank are
united, the oldest legitimate name or (for taxa below the rank of
genus) the oldest legitimate epithet is retained . . .”” As such,
Acanthopanax (Miq.) Harms is illegitimate, and so is A. sen-
ticosus (Rupr. & Maxim.) Harms.

Finally, it is our opinion that only critical studies covering
the entire taxonomic and geographic ranges of Eleutherococ-
cus and Acanthopanax will enable one to judge correctly, if
union of these two taxa is warranted.

Eleutherococcus senticosus (Rupr. & Maxim.)
Maxim., Mém. Présent. Acad. Imper. Sci.
St.-Pétersb. Ser. 6c, 9: 132. 1859; Nakai, J.
Arnold Arbor. 5: 10. 1924; Poyarkova in
Shishkin, Fl. USSR 16: 17. 1973.

Hedera senticosa Rupr. & Maxim., Bull. Phys.-
Math. Acad. Imper. Sci. St.-Petersb. 15: 134.
1857.

Acanthopanax senticosus (Rupr. & Maxim.)
Harmsin Engler & Prantl, Naturl. Pflz. Fam.
3(8): 50. 1898; Li, Sargentia 2: 71. 1942.

Acanthopanax eleutherococcus Makino, Bot.
Mag. Tokyo 12: 19. 1898. (Fide Nakai, 1924;
Li, 1942; Poyarkova, 1973).

Eleutherococcus koreanus Nakai, Fl. Sylyv.
Koreana 15:32. 1926. (Fide Nakai, 1924; Li,
1942: Poyarkova, 1973).

342

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Dardymov, I.V. 1976. Ginseng and Eleutherococcus, Ist ed. Nauka Publish-
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Rehder, A. 1922. Acanthopanax. /n Bailey, L.H., The Standard Cyclopedia
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Schneider, C.K. 1912. Illustriertes Handbuch der Laubholzkunde, Band 2, p.
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343