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BOTANICAL MUSEUM 
LEAFLETS 
HARVARD UNIVERSITY 


PRINTED AND PUBLISHED AT THE 
BOTANICAL MUSEUM 
CAMBRIDGE, MASSACHUSETTS 


BOTANICAL MUSEUM LEAFLETS 
HARVARD UNIVERSITY 


VOLUME XXIV 


BOTANICAL MUSEUM 
CAMBRIDGE, MASSACHUSETTS 
1974-1976 


TABLE OF CONTENTS 


NUMBER | (July 31, 1974) 


Archeological Evidence for Snuffing in 
Prehispanic Mexico 
By Pati PONS coyeenenndecnacna den pes teks i 


NUMBER 2 (October 25, 1974) 


A Study of Systematic Wood Anatomy in Cannabis 
By LoRAN C. ANDERSON .........0cc cece ceeeeces 29 


Two New Brazilian Species of Brunfelsia 
By TIMOTHY C. PLOWMAN ...............-0eeee ee OU 


NUMBER 3 (January 10, 1975) 


Archeological Maize from Northern Chile 
By PauL C. MANGELSDORF AND 
CORDON Foil AW een sa ong te acaanaes 49 


NuMBER 4 (June 16, 1975) 


The Contribution of Linnaeus to Orchidology 
By WILLIAM T SPEARN 9 co-aton4 nd bteedegeus ideas 65 


NUMBER 5 (June 27, 1975) 


Ethnobotany of the Western Tarahumara of Chihuahua, 
Mexico, I. Notes on the Genus Agave 
By ROBERT A. BYE, JR., DON BURGESS, AND 
ALBING MARES TRIAS. vcuw ores bah5 or tah dad vex aon 85 


NuMBER 6 (October 31, 1975) 


Nutritional Value of Coca 
By JAMES A. DUKE, DaviD AULIK AND 
TIMOTHY PLOWMAN ...........cccccccccccccesves LES 


De Plantis Toxicariis e Mundo Novo Tropicale 
Commentationes XIII. Notes on Poisonous or 
Medicinal Malpighiaceous Species of the Amazon 
By RICHARD EVANS SCHULTES ........- +e ee eeeees 121 


NUMBER 7 (December 12, 1975) 


Studies in the Genus Oncidium. I 


DSU Fe SEAGY. tavuep su iaveeeseereeke napa ee 133 


NuMBER 8 (April 9, 1976) 


Plantae Colombianae XIX. E Partibus Amazonicis 
Witotorum Plantae Fructuariae Sativae Novae 
By RICHARD EVANS SCHULTES «2. ...00. 0008s eeees 193 


Pachytene Chromosome Morphology and 
Its Bearing on Interspecific and Intergeneric 
Relationships of Coix 
By J. VENKATESWARLU, R.S.K. CHAGANTI AND 
PANUCANTI NN. RAG: beeee se iisdeeias ees be esees 205 


New Chichuahuan Umbelliferae 
By LINCOLN CONSTANCE AND ROBERT A. BYE, JR... 229 


Chromosome Numbers in Chihuahuan Umbelliferae 
By LINCOLN CONSTANCE, TSAN [ANG CHUANG AND 
ROPER T Ba PI, ga sc chara ed pee yes Chee Ses 24] 


NuMBER 9 (June 30, 1976) 


The Genus Oeceoclades Lindl. 
By LESLIE A. GARAY AND PETER TAYLOR ........- 249 


NuMBER LO (November 30, 1976) 
Manicaria saccifera and Its Cultural Significance 
Among the Warao Indians of Venezuela 
By JOHANNES WILBERT sxi 544s ccee Ses sqm wees 2795 


[ vi | 


INDEX OF ILLUSTRATIONS 


PLATE 
Agave americana var. expansa ............0.000- XVII 
Agave bovicornuta .............cccccceecceeees XVIII 
Agave multifilifera ..............0.. 0 ccc eee ee XXII 
AGAVE DACINCS. oni uGale tied eae ed bbe de heeds e ae nns XIX 
Agave polianthiflora ............ 0... cee eee eee XXII 
Agave SATEVEl cad iiwine ii ers ¥e% beeen beannei XX 
Agave Vilmoriniana ................c cece eeeeeee XXIII 
Agave WOCOMIAML kana s iss oyeecusatiah cae codunews XXI 
Brimrersia Dil0Ga. s+ peeve nso el ecek een vor8 XIII, XIV 
Brunfelsia Martiana ............0cc cece ee eees XI, XII 
Cannabis wood sections .......... 0.0.0. ccc cee cence X 
Coix 
Ideograms of pachytene chromosomes .......... LX 
Five pachytene chromosomes of C. aquatica ... LVI 
Pachytene nucleus of C. aquatica .............. LV 
Pachytene nucleus of microsporocyte 
GE CG. ZiPanted  .4scad bc Sods eeu Veedei ces edcana LIX 
Pachytene nucleus of C. Lachryma-Jobi ...... LVIII 
Pachytene nucleus of microsporocyte of 
C. Lachryma-Jobi ............ 0. cee cee euees LVII 
Corn cobs, late prehispanic ...............0 00000. XV 
Corn cobs, dated at ca. A.D. 100 and 1250 ........ XV 
Donnellsmithia silvicola .............. 002.0000 LXI 
PYyneiiin Gentry) s+ swe cenieed owihyie ee cees LXIV 
Heteropteris macrostachya 444. é.s ised ivaeia sae MLV 
Pie@terapreris Fiala coh<eseehecie cay en se cchanncelVy 
PRinA@a APADOTOUSIS +. ie n:kst nau ee dak ad os od 
[niga @norerana.- 4.66222 <40y dx cic cde wedavnwacadvs LIV 
Macoubea guianensis ......... 00.0 cece eee eee ase LII 
Macoubea witotorum ............c00 cece eee XLIX, L 
Macoubea wititorum, fruits of ..............0-0 000s Li 
Manicaria saccifera 
CAOWINe ti 1Otest 64 ees 4aeabaidted sein escent’ LXVI 
Details of inflorescence ...... LEAVE, bas VL LA 
Thatch on Warao houses ....... LAX, DAS, LAXH 


Leaves used for sails on canoes 


LXIII 


PER any eee See ee ae, 


AUIS saqd2au Swneee ter eres ea ogee he aes LXXIV 
Hats made from spathes .... LXXV, LXXVI, LXAXVII 
Fruiting spathe and infructescence ....... LXNXVITI 
TOP WRU Cd OI SOR, ne siyss oto eved Soho ass LXXIX 


Steps in the processing of sago 


Map of Karaparana-Igaraparana region of 


Colombian Amazon 
Mascagnia glandulifera 
Mauritia flexuosa trough 
Mochica IV terracotta vessel 


Oncidium 
Oncidium 
Oncidium 
Oncidium 


Pome nr me SAN er eee Sar ar ee ee 


Ce 


LXNXXIT, LXXXITI 
ea ee es eee bene s IX 
... XXXVI, XLVIII 


aequinoctiale (in color) 


alticola (in color) ......XXXV, XLV, XLVII 
ANCIPERUGM fo naae en ce siadeneesis ges oa 
APUAVENSE  dndeneans chawdedn ee ieee 


Oncidium chimborazoénse (in color) .XXXVII, XLVII 
Oncidium cucullatum (in color) ......XXITX, XXXVITI 
Oncidium cucullatum var. dolabratum ..........XXDX 
Oncidium cucullatum var. macrochilum ........ XIX 
Oncidium Dayanum (in color) .... XXXI, XLII, XLIV 
Oncidium erosilabium (in color) ......XXXVI, XLVII 
Oncidium Kennedyi (in color) ................ XXXIX 
Oncidium mimeticum (in color) XXXII, XL, XLI, XLII 
Oncidium nubigenum (in color) ........ XXXIV, XLVI 
Oncidium olivaceum (in color) ..... XXVIII, XXXVITI 
Oncidium olivaceum var. giganteum ......... XXVITI 
Oncidium Phalaenopsis (in color) .............. XLII 
Oncidium rhodostictum .............. XXX, XXXVITI 
Oncidium sanguinolentum (in color) ..... XXXII, XLI 
Oncidium spathulatum (in color) ........ XXXII, XLIV 
Oncidium tarquiense (in color) ..........XXXV, XLV 
Oncidium tripterygium (in color) ............... XLII 
Oncidium tunguraguense (in color) .. XXXVIT, XLVIII 
UTIs. “nech ees ac oecae ewe eens eae bad oe a I - VIII 
TASCA OCG 2nbese cian ceeeee eee has es owes LXII 
TP etisC ie PAPAIN. enact ees erred Lae eae teas LXIII 
Petraptens Muctonat®: s425004 441-44 50544es rene VEL 
Tetrapteris styloptera .......-.-.-+-000++0+.+. MXVII 
Triorchis sera (Orchis morio) .............002+++. V1 
Umbellifereae, karyotypes of Chihuahuan ....... LXV 


[ vill | 


AECEOCLADES 253 

AEROBION 
maculatum 266 

AGAVE 85 
americana 

var. expansa, 

bovicornuta 94 
multifilifera 97 
nomenclature of Western 
Tarahumara LOL 
pacifica 95 
polianthiflora 97 
Shrevei 96 
Vilmoriniana 97 
species of Western 


93 


Tarahumara LOO 
Wocomahi 96 
ahue 91 
AMANITA 
muscaria 22 
ANADENANTHERA 
colubrina 22 
ANGRAECUM 
maculatum 253,254,266 
monophyllum 268 
ARRACACIA 245 
atonali 88 
balila 89 
BANISTERIOPSIS 
Caapi 3,122 
Martiniana 
var. laevis 123 
bee-ra-ree-a-ma_ 130 
BRUNFELSIA 
pilosa 42 
Martiana 
CANNABIS 
indica 29 
sativa 29 
wood anatomy 29,31 
chahui 93 
cobisi 89 


37 


INDEX 


[ioe 4 


coca 113 
117 


insecticidal residues 117 


alkaloids in leaves 


nutritional comparison 119 
nutritional value 
COIX 
aquatica 207 
chromosome numbers 205 
gigantea 210 
Lachryma-Jobi 210 
COPERNICIA 
cerifera 277 
curanderos 3 
CYMBIDIUM 
calcaratum 261 
DONNELLSMITHIA 241 
silvicola 226 
eé-taw-gaw 126 
emukohoko 300 
EPIDENDRUM 
connivens 267 
ERYNGIUM 245 
Gentryi 235 
ERYTHROXYLUM 
coca 113 
EULOPHIA 254 
alismatophylla 258 
ambongensis 268 
atrovirens 260 
Bierleri 271 
calcarata 261 
cordylinophylla 261 
Decaryana 262 
dissimilis 264 
Elliotii 271 
gracillima 263 
Ledienii 266 
lanceata 263 
latifolia 264 
lokobensis 261 
lonchophylla 264 
Lubbersiana 265 


ies 


Mackenii 265 
maculata 266 
Milbraedii 270 
pandurata 268 
paniculata 261 
petiolata 269 
quadriloba 269 
Saundersiana 270 
Schlechteri 258 


sclerophylla 271 
seychellarum 272 
spathulifera 272 


tainioides 264 
ugandae 273 
,ULOPHIDIUM 253 
alismatophyllum 258 
ambongense 258 
analamerense 259 
analavelense 259 
angustifolium 259 
ssp. diphyllum 259 
boinense 260 
cordylinophyllum 261 
Decaryanum 262 
dissimile 264 
gracillimum 263 
latifolium 264 
Ledienii 266 
lobense 262 
lonchophyllum 264 
Lubbersianum 265 
Mackenii 265 
maculatum 266 
var. pterocarpum 
monophyllum 268 
nyassanum 267 
paniculatum 261 
panduratum 268 
Perrierit 268 
petiolatum 269 
quadrilobum 269 
Rauhii 270 
roseovariegatum 270 
Saundersianum 270 
sclerophyllum 271 


267 


seychellarum 272 
spathuliferum 272 
tainioides 264 
Warneckeanum 266 
zanzibaricum 273 
galime 92 
GEODORUM 
pictum 266 
GRAMINEAE 205 
GRAPHORCHIS 
260 
maculata 266 
Saundersiana 270 
vue huali 89,90 
gusime 92 


atrovirens 


hallucinogens 1 
henequén 85 
HETEROPTERIS 
macrostachya 124 
riparia 123 
suberosa 124 
HIRAEA 
apaporiensis 124 
Schultesii 124 
ho-ree-a-mee-see 


234 


130 

huve 

ILEX 
Guayusa 3 
paraguariensis 3 

ime 92 

ime juice 89 

INGA 


chorrerana 202 


JUSTICIA 4 


LEOCHILUS 
sanguinolentus 152 
LIGUSTICUM 245 
LIMODORUM 
maculatum 266 
LISSOCHILUS 254 
ambongensis 268 
analamerensis 259 
analavelensis 259 
barombensis 271 


boinensis 260 


cordylinophyllus 261 
Decaryanus 262 
Elliotii 271 
gracillimus 263 
Hebdingianus 263 
lokobensis 262 
lonchophyllus 264 
panduratus 268 
paniculatus 261 
petiolatus 269 
quadrilobus 269 
Schlechteri 258 
spathulifer 272 

litos 5 

LOPHOPHORA 
Williamsii 3 

MACOUBEA 
guianensis 198 
witotorum 193 

maguey 85 

maize 
archeological 49 
Capio Chico 61 
Chutucuna Chico 61 
cobs from site A.D. 105 51 
Confite Iqueno 53 
Confite Morocho 57 
Confite Puntiagudo 57 
Polulo 53 
Rabo de Zorro 57 
sting cobs 57 

Malpighiaceous Amazonian 
species 
medicinal 121 
poisonous 121 

MANICARIA 

saccifera 275 
composition of starch 319 
ethnobotanical lore of 319 
ethnomedicine of 303 

MASCAGNIA 
glandulifera 126 

masiavari 230 

maté 3 

mee-see-gaw 123 


mesagoli 89 

mescal bread 89 

MEZIA 
includens 126— 

NICOTIANA 2 

no-na-mee-koo-ma_ 130 

OECEOCLADES 249,253 
alismatophylla 258 
ambongensis 258 
analamerensis 259 
analavelensis 259 
angustifolia 259 
atrovirens 260 
boinensis 260 
calcarata 261 
cordylinophylla 261 
Decaryana 262 
gracillima 262 
Hebdingiana 263 
key to species 254 
lanceata 263 
latifolia 264 
lonchophylla 264 
Lubbersiana 265 
Mackenii 265 
maculata 266 

var. pterocarpa 267 

monophylla 267 
pandurata 268 
Perrieri 268 
petiolata 269 
quadriloba 269 
Rauhii 270 
roseovariegata 270 
Saundersiana 270 
sclerophylla 271 
seychellarum 
spathulifera 2 
ugandae 273 
zanzibarica 273 

ojcome 92 

ONCIDIUM 133 
aequinoctiale 162 
alticola 159 
andigenum 151 


a oe | 
~l 
aw 


azuayense 156 
chimborazoénse 164 
cucullatum 137,144 
subvar. andigenum 151 
subvar. spathulatum 148 
var. andigenum 151 
var. Chestertoni 148 
var. Dayanum 150 
dolabratum 144 
flavidum 156 
maculosum 146 
microchilum 146 
nubigenum 158 
olivaceum 14] 
Phalaenopsis 150 
purpurascens 146 
sanguinolentum 152 
spathulatum 148 
Dayanum 150 
erosilabium 162 
Kennedyi 142 
mimeticum 152 
var. flavidum 156 
nubigenum 158 
var. spathulatum 148 
olivaceum 141 
var. andigenum 15] 
var. flavidum 156 
var. giganteum 141 
var. Lawrenceanum 
var. maculosum 146 
var. nubigenum 158 
var. Phalaenopsis 150 
var. sanguinolentum 152 
Phalaenopsis 150 


var. 
var. 
var. 
var. 
var. 
var, 
cdr. 
var. 
vdr. 
cdr. 


142 


var. Brandtiae 150 

var. excellens 148 
rhodostictum 146 
sanguinolentum 152 
spathulatum 148 
tarquiense 160 
tripterygium 151] 
tunguraguense 166 


Orchids of Linnaeus 65 
o tosa 92 


[ xii | 


palm, Temiche 286 
pevote 3 
pipes, nose 14 
Piptadenia 3 
pod corn 57 
popcorn, Peruvian 57 
pottery artifacts, Oaxaca 16 
PRIONOSCIADIUM 245 
PSEUDOCYMOPTERUS 245 
PSYCHOTRIA 4 
pueblo duck mythology 18 
RIVEA 
corymbosa 3 
rolobusi (Bovardia sp.) 87 
SACCOLABIUM 253 
sapogenin 91 
sa puli 92 
sarabatucu. 124 
sepe 235 
shamans and shamanism 
sisal 85 
smilagenin 91 
snuff 1 
hallucinogenic 4 
snuffers and snuffing 1,6 
Ixtlan 10 
Virola 4 
SOPHORA 
secundiflora 24 
SORGHUM 
intrans 218 
purpureo-sericeum 218 
starch, palm 275 
stupifving fish 91 
sugui (tesguino) 88 
SYAGRUS 
Romanzoffianum 277 
takwatsi 22 
tamatsi Maxa Kwaxi 
tarahumaras 89 
TAUSCHIA 245 
bicolor 229 
tarahumara 232 
TETRAPTERIS 
mucronata 128 


90 


ae 


20) 


silvatica 128 wee-po-awk 130 
styloptera 128 willka 22 
tortillas 90 Xochipala art 12,14 
UMBELLIFERAE 225 vahuhi aru 279 
chromosome numbers 241 yvawihi aukwaha 300 
usabi (Prunus Gentryi) 87 yaje 123 
vilca 22 ve-aing 124 


[ xiii | 


a i ae 


oe 


~ BOTANICAL MUSEUM LEAFLETS 
“a HARVARD UNIVERSITY 


CampripGr, Massacnuserrs, Jury 31, 1974 Vor. 24, No. 1 


ARCHAEOLOGICAL EVIDENCE 
FOR SNUFFING IN PREHISPANIC MEXICO 
BY 
Perer T. Furst 


The various hallucinogenic or psychoactive plants the 
extensive religious and divinatory use of which by Mexi- 
‘an Indians both fascinated and appalled the Spanish 
colonial clergy of the sixteenth and seventeenth centu- 
ries were smoked, chewed, sucked, brewed or macerated 
into beverages and otherwise ingested in liquid, solid or 
incinerated form. ‘Tobacco, most commonly smoked, 
was also ground into a fine green powder that was not 
taken internally but was rather applied externally to the 
patient's body in shamanistie curing practices. 

All these customary uses of ‘‘mind-altering’’ sub- 
stances are well described for both prehispanic and post- 
Conquest Mexico by such writers as Sahagin, Duran, 
Hernandez and, somewhat later, Jacinto de la Serna and 
Ruiz de Alarcon. In contrast, there is no mention what- 
ever of hallucinogenic snuffs, taken through tubes or 
through ‘‘nose pipes’’, a common practice in the West 
Indies and in Central and South America. Since these 
and other chroniclers of indigenous practices and beliefs 
were usually careful observers, and since the Church was 
engaged in a vigorous—albeit ultimately unsuccessful— 
campaign to discover and suppress the indigenous use of 
intoxicants of all forms, we must assume that, trade and 


[1] 


ECONOMIC BOTANY * \RY. 


las Fa 
Le 


other contacts with Caribbean and Central American 
snufE~using cultures notwithstanding, the Indians of 
Late Post-Classic Mexico seem not to have assimilated 
these practices into their own extensive complex of 
ritual intoxicants. 

Nevertheless, there is a growing corpus of data, in the 
form of archaeological art, to suggest that snuffing was 
once known and. practiced in several parts of Meso- 
america as early as 1500-1200 B.C. and at least as late 
as the first centuries A.D. Before we examine some of 
the evidence, we need briefly to consider the problem of 
potential indigenous Mexican sources for hallucinogenic 
snuffs. 

Thanks to prodigious research in the field, the labora- 
tory and the historical sources, especially by such inves- 
tigators as Richard Evans Schultes, S. Henry Wassén, 
Siri von Reis Altschul, and Bo Holmstedt, the various 
kinds of South American snuffS are botanically and 
chemically rather well understood. For Mexico, how- 
ever, the botanical data are inadequate, and chemical in- 
formation is wholly or largely lacking. Nevertheless, 
there are indications of the direction that future research 
might fruitfully take. 

First, one cannot rule out one or more species of Nico- 
tiana. These native tobaccos have a much greater nicotine 
content than do the hybrid species from which cigarette 
or pipe tobaccos are made. Wilbert (1972: 55-73) re- 
cently documented the use of tobacco as the sole psycho- 
tomimetic employed by the shamans of the Warao In- 
dians of Venezuela, who smoke themselves into ecstatic 
trance states that are phenomenologically indistinguish- 
able from those elsewhere triggered with such botanical 
hallucinogens as ayahuasca, the sacred mushrooms, 
morning-glory seeds, or cfnadenanthera and Virola 
snuffs. Several species of Nicotiana are in tact employed 


[2] 


for psychotomimetic snuff in South America, either 
alone or in combination with other psychoactive plants. 
The shamans of some indigenous cultures—e.g., the 
‘Tacana of Bolivia—use pulverized tobacco as a magical 
repellent against hostile demons (Hissink and Hahn, 
1961). Furthermore, it is not impossible that the use of 
tobacco powder as a magical external medicine by Mexi- 
‘an curanderos had its ultimate origin in an earlier use 
of powdered tobacco as snuff. 

Secondly, there appears to be no reason why some of 
the better known Mesoamerican plant hallucinogens 
should not be as psychotomimetically effective when 
taken as snuff through the nasal membranes as when as- 
similated through the stomach. By way of illustration, 
I am informed by Dr. Schultes that, in South America, 
the bark of Banisteriopsis Caapi, which is usually macer- 
ated or brewed into the potent hallucinogenic beverage 
known by such names as yajé, ayahuasca, etc., is re- 
putedly sometimes pulverized and inhaled as snuff. Even 
Ilex Guayusa, a caffeine-rich holly widely utilized as a 
stimulating tea, along with its sister species, e.g. mateé, 
Ilex paraguariensis), has served as snuff, at least in pre- 
hispanic highland Bolivia, where Ilex snuff and snufhing 
paraphernalia were recently discovered in a Tihuanacoid 
shaman’s grave, dated ca. A.D. 500. The shaman’s kit 
also included clysters, suggesting that the same plant 
might even have been employed for stimulating enemas 
(Schultes, 1972b). 

Whether or not peyote (Lophophora Williamsi), olo- 
liuhqui (Rivea corymbosa) or other hallucinogens native 
to Mesoamerica were ever used in the form of snuff, 
there exist extensive Mexican populations of shrubs and 
trees of the Leguminosae that should be investigated for 
possible psychoactive properties. Included are two Mexi- 
can species of Piptadenia, a genus closely related to the 


[ 3 ] 


psychoactive genus Anadenanthera of South America, 
which may well possess the same or similar psychotomi- 
metic constituents. Two of these Mexican species are 
Piptadenia flava, found also in Central America and Co- 
lombia, and Piptadenia constricta. Both are found along 
the Pacific coast, from Sinaloa and Jalisco in the north 
to Guerrero in the south. To my knowledge, neither 
have been tested for hallucinogenic alkaloids. There are 
in addition more than sixty species each of the allied 
genera Mimosa and Acacia in Mexico, and some of these 
may, like certain South American species, contain psy- 
choactive principles. This might be found to apply es- 
pecially to those species credited with sacred, magical, 
or ‘“‘dangerous qualities by local Indians or rural mez- 
tizos and should be chemically studied from the point of 
view of possible hallucinogenic alkaloids. 

Finally, there appear two other possible candidates as 
potential sources of hallucinogenic snuff in southeastern 
Mexico, both with significant South American. ties. 
These are species of Psychotria and Justicia. The former 
is a well known additive in hallucinogenic potions pre- 
pared basically from the Banisteriopsis Caapt vine in 
Ecuador, Colombia, Peru and Brazil, while the latter is 
added to /rola snuff or is said even to be employed 
alone as a source of psychotomimetic snuff (Schultes, 
1972a: 45-46, 52). These possibilities emerge from a 
comment by Wassén (1972: 37-88) on a suggestion of 
mine (1968: 160-164) that snuffing might have been 
practiced by the Gulf Coast Olmec. In support of this 
comment, Wassén cites the following excerpt from a let- 
ter to him by Schultes, dated February, 1969: 


We are finding so many plants with tryptamines—the active 
principle of many of the snuffs of South America— that it is very 
possible that in the Mexican Gulf Coast area the Indians could 
have found a plant which, prepared in the form of a snuff, could 
intoxicate as does the snuff of the Waikas. One of these is Psy- 


7 


chotria, a species of which in South America has now been found 
to have N,N-dimethyltryptamine. 

Psychotria occurs up as far as Vera Cruz and it is possible that 
other species have this principle. Furthermore, Holmstedt be- 
lieves that he has found this same chemical in our species of 
Justicia which is added to Viro/a snuff by the Waikas. Other 
species of Justicia occur as far north as Vera Cruz and may pos- 
sibly also have this chemical constituent. 


While the botanical sources for hallucinogenic snuff 
in Mesoamerica must, for the present, remain conjectur- 
al, the evidence for snuffing in archaeological art is, as 
we shall see, beyond question. Moreover, on the earliest 
level of the proposed Mesoamerican snuffiing complex— 
that is, the Early to’ Middle Formative—the evidence 
points persuasively southward, at least to Central Ameri- 
‘a, if not actually to northwestern South America. 

We owe much of our knowledge of Central and South 
American snufting paraphernalia—prehistoric as well as 
recent—to Wassén’s several studies, and I would here 
like to acknowledge my own debt to our Swedish col- 
league in this area of research. It was a paper by Was- 
sén, published in 1967, that first set me on the track of 
a possible snuffing complex in Mexico. Specifically, my 
attention was drawn to the so called Brazilian “tos, small 
effigy stone carvings, usually bird-like, with carved, shal- 
low, oval or circular depressions that made them appear 
like receptacles. A number of these were found in the 
last century in the shell middens of Santa Catarina, 
Brazil. Wassén thought it likely that these bird-efhgy 
litos might have served as tablets for hallucinogenic 
snuff, rather like the archaeological wooden snuff tablets 
found in the Chilean and Peruvian desert, or more re- 
cent snuff tablets from Amazonia. 

Subsequently, I raised the question of the use of hal- 
lucinogens by the Olmec, suggesting that the well known 
jade artifacts called ‘*spoons’’, might, like the Brazilian 


[ 5 | 


litos, have served as snuff tablets (Furst, 1968: 162). At 


least some of the Olmec ‘‘spoons’’ seemed to represent 
long-tailed birds in flight, seen in profile. In any event, 
like some South American snuff tablets, certain Olmec 
jade spoons are decorated with bird-jaguar motifs, a com- 
mon symbolic theme in South American ritual intoxi- 
‘ation. 

At the time, this was still highly speculative. No 
direct evidence existed to show that the Olmec had used 
snuff or other hallucinogens; for that matter, there was 
no proof that the ritual use of psychoactive substances 
in Mesoamerica was any older than the oldest of the so- 
‘alled mushroom stones, 1.e., from the end of the Mid- 
dle to the Late Formative. All that could be said was 
that it would be surprising if the Olmec had used no 
hallucinogens, considering what was already known of 
the antiquity and wide distribution of the hallucinogenic 
phenomenon in the New World. ‘To mention only snuff 
ing: the earliest known archaeological snuffing imple- 
ments are a whalebone tablet and associated birdbone 
snuffing tube which Junius Bird of the American Mu- 
seum of Natural History excavated at Huaca Prieta, 
Peru. These are dated at ca. 1500-1700 B.C. The evi- 
dence was thus conclusive fora time depth of some 8,500 
years for the use of, hallucinogenic snuffs in South 
America. 

The first evidence that snuffing was in fact known at 
one time also in Mesoamerica came to my attention in 
the form of a hollow, redware effigy figurine from Co- 
lima, representing a seated man with a horn on his head 
and a small, gourd-shaped nose pipe held to one nostril 
(Plate 1). Subsequently, [ was to come across a second, 
considerably larger, Colima effigy (Plate I), of burnished 
brown clay, sculpturally far more sophisticated, again 
depicting a manin the act of snuffing from a gourd- 


[6 ] 


PLATE I 


Small, hollow terracotta efigy of seated man, holding a gourd- 


shaped snufling pipe to his nose. Colima, shaft-and-chamber tomb 
~ly 


phase, ca. 100 B.C.-—A.D. 200. Anon. private collection. H. 75 


PLATE II 


Effigy figurine of burnished brown clay depicted in the act of in- 
haling snuff from a bottle gourd-shaped nose pipe. Colima, shaft- 
and-chamber tomb phase, ca. 100 B.C.—-A.D. 200. Kurt Stavenhagen 
Collection, Mexico City. H. 11”. 


shaped nose pipe. Both of these figurines belong to the 
larger West Mexican shaft-and-chamber tomb art com- 
plex and can, therefore, be dated between 100 B.C. and 
A.D. 100-200. 

In addition to the effigies, we were able to identify, 
in several collections, a number of pottery snuffers or 
nose pipes from West Mexico that closely resemble the 
well known Costa Rican snuffing pipes illustrated by 
Weassén in several publications. [specially interesting is 
a red-slipped snuffer with bifurcated stems, one for each 
nostril, from the Ixtlin del Rio area of southern Nayarit 
(Plate IIL). The Ixtlin snuffer is actually a convention- 
alized bird effigy, with nubs at the side of the bowl to 
indicate wings, and a projection at the front for the head 
or beak. Such abbreviated bird symbolism is common on 
Costa Rican pottery snuffers as well. That this is hardly 
fortuitous was recognized by Wassén: birds and_ bird 
spirits are widely connected with the ecstatic trance ex- 
perience and with shamanism. 

For atime, these West Mexican specimens seemed to 
be all that there was. Snuffing, therefore, appeared to be 
an isolated phenomenon in time and space, associated with 
the shaft-and-chamber tomb cultures of the West Coast. 
Their funerary art indicates that these same cultures also 
employed the peyote cactus and, possibly, mushrooms. 
The close similarity of the West Coast pottery snuffers 
to those of Central America, and their restricted distri- 
bution close to the Pacific coast, suggested a somewhat 
short-lived trait, introduced possibly from a southerly 
source, that eventually failed to take hold alongside es- 
tablished cults involving such well known indigenous 
Mexican hallucinogens as peyote and the sacred mush- 
rooms. 

However, West Coast snuffing was not to remain the 
isolated and short-lived phenomenon that it appeared at 


/ 10 | 


Puatre Ill 


Bifurcated bird effigy snuffer from Ixtlan del Rio, Nayarit, shaft- 
and-chamber tomb phase, ca. 100 B.C.-A.D. 200-300. Coll. Mr. 
and Mrs. William Kaplan, New York. L. 24”. 


first. New evidence has come to light in the form of effigy 
and undecorated snuffing pipes from the Karly to Middle 
Formative, from Nochipala, Guerrero. The dating of 
XNochipala is still somewhat uncertain: there have been 
suggestions that it represents the very ‘“‘origin’’ of Olmec 
art, predating the San Lorenzo Phase in Veracruz (Gay 
1972). However, the reported and confirmed associations 
of the extraordinarily sophisticated and sometimes as- 
tonishingly naturalistic figurines from Nochipala with 
typically Olmec incised bowls and other Olmee artifacts 
characteristic of the Late Karly to Karly Middle Forma- 


6 


tive, including *“*spoons’’ and beads of blue-green trans- 
lucent jade, suggests dates equivalent to San Lorenzo 
and contemporaneous sites in Morelos and elsewhere in 
Central Mexico—i.e., between 1800 and 1000 B.C. On 
the other hand, a recent series of thermoluminescence 
tests tend to support an earlier date at least for the be- 
ginning of realistic Nochipala art, possibly as early as 
1500-1600 B.C. (Robert Stroessner, pers. comm. ). 
The uncontrolled looting of Nochipala, with its re- 
markable assemblage of some of the finest Formative 
ceramic figurines to be found anywhere in the New 
World, is a scientific tragedy of major proportions. We 
‘an only guess at the evidence that has forever been lost ; 
nevertheless, it has been possible to study and verify the 
authenticity of a number of Nochipala pieces now in 
private hands or museum collections (interestingly 
enough, although the accidental discovery of the Nochi- 
pala site by local farmers dates from the mid-nineteen 
sixties, a typical Nochipala figurine has been in the ex- 
tensive pre-Columbian collection of the Peabody Mu- 
seum of Harvard University for more than seventy 
years). Among recently discovered Nochipala artifacts 
are several unmistakable snuflfing instruments or ‘‘nose 
pipes’, dating far earlier than those of the West Coast 


[ 12 | 


Piatt LV 


Terracotta bowl snuffer, Xochipala, Guerrero, Early Formative, 
ca. 1500-1200 B.C. Anon. private collection. L. 24”. 


[ 13 | 


shatt-and-chamber tomb phase and approaching in an- 
tiquity the earliest South American paraphernalia found 
on the Peruvian coast. 

The first of these to be examined and identified as a 
nose pipe was a small, round, undecorated bowl with a 
horizontal perforated stem (Plate IV). If one compares 
this pottery snuffer with examples from Central America, 
it is clear that, except for its characteristic local paste 
and the lime encrustation typical of ceramics from the 
XNochipala burials, the little Mexican snuffing pipe is 
virtually identical to similar instruments from Guana- 
‘caste or Linea Vieja, Costa Rica (Wassén, 1965: 25). 
As in the case of the Nayarit snuffer, it is difficult not 
to postulate a genetic connection between them, al- 
though the known Central American pottery snuffers, 
and also that from Nayarit, are appreciably later than 
this Early Formative pipe. 

A second Nochipala nose pipe which | was able to 
study in detail is much more complex (Plate V). It is 
an effigy pipe, measuring 4?” in length, representing a 
human figure on its back, with knees drawn up—a posi- 
tion somewhat resembling the post-Classic *“*Chacmool”’ 
stone sculptures. On the basis of the wrap-around loin- 
cloth, the figure can be identified as male. In a recent 
museum catalogue, the piece was erroneously described 
as an efigy bow] in the form of a kneeling person (Gay 
1972). But that would place the nosepiece at the top 
and the bowl opening facing vertically toward the front, 
which seems hardly likely. Once the piece is recognized 
for what it is—a nose pipe used for snuffing—the location 
and inclination of the nosepiece alone dictate a supine 
position for the figurine, as does the opening of the bow] 
itself. Indeed, in handling the piece, its real purpose 
suggests itself almost spontaneously. 

While it is certainly the finest example known to me, 


[ 14 J 


[ ¢r ] 


Ethgy snuffer representing a man lying on his back. The burnished nosepiece emerges 
hornlike from the topof his head. From Xochipala, Early Formative, ca. 1500-1200 B.C. 
Anon. private collection. L. 4}. 


ALWIG 


A 


this effigy pipe is by no means unique in form. Several 
similar nose pipes have come to light, including one 
(Plate VI) of aman with animal characteristics lying on 
his stomach, with the bowl in the back and a horn-like 
nosepiece on top of the head. Here again the sex of the 
efligy is male; indeed, there are indications of a phallus 
on the underside. 

On the basis of the evidence, then, we can postulate a 
snufling complex of appreciable duration and antiquity 
along the west coast of Mexico, with the earliest evi- 
dence dating to 1500-1200 B.C., and the latest approxi- 
mately to the beginning of the Christian era. 

A recent re-examination of early pottery artifacts from 
Oaxaca as well as from Central Mexico shows, however, 
that divine inebriation with psychotropic snuff was not 
limited to the Guerrero Formative or the shaft-and- 
chamber tomb phase of coastal northwestern Meso- 
america. | have only just begun checking through col- 
lections and the literature on Monte Alban ceramics, but 
already it appears that the evidence for snufling from the 
Late Formative at least into the Karly Classic is sub- 
stantial. Thus far, we have been able to identify more 
than a dozen spouted ‘‘miniature effigy vessels’*, includ- 
ing a group in the Museo Frissell de Arte Zapoteca in 
Mitla, Oaxaca, as probable nose pipes, dating from Monte 
Alban I and II. Some of these appear very similar in 
construction, if not in style and paste, to those from 
XNochipala. In addition, I have located at least one 
probable Early to Middle Formative animal effigy nose 
pipe, in the form of a turtle, from Tlatilco, in the col- 
lections of the Museum of Ethnology in Vienna, Austria. 
This relates stylistically to black effigy ceramics of Olmec 
derivation or origin from Tlatilco and Las Bocas, Puebla. 

One interesting little polished black snuffing pipe, 
possibly transitional from Monte Alban I to IT (i.e. ca. 


[ 16 | 


PLATE VI 


Effigy bowl snuffer in the form of a person with human head and 
animal-like body, from Xochipala, Guerrero, Karly Formative, 1500-— 
1200 B.C. Asin Plate V, the nosepiece is on top of the head. Anon. 
private collection. L. 4’. 


200 B.C.), appears to symbolize transformation, in this 
instance from human into a duck-like bird with rounded 
body and flipper-like feet (Plate VII). The frontal half 
is human, with hands held palms together to the chin; 
the rest of the body is that of a duck. A conical per- 
forated horn on the head forms the nosepiece, as in some 
of the effigy pipes from Nochipala. 

Space limitations preclude detailed discussion of duck 
symbolism, but it should be noted that chimereal or 
anthropomorphic ducks are not uncommon in_prehis- 
panic art, especially en the west coast. Ducks are present 
also in the art of Tlatilco and other Karly to Middle 
Formative sites. A study of Pueblo duck mythology and 
behets about the duck as supernatural among the Cor: 
and Huichol of West Mexico may throw some light on 
the problem; a Duck Person is a prominent figure in 
Huichol origin myths, as it is also among the Zuni and 
other Southwestern Indians; ducks seem to be messen- 
gers of the gods or else a form that the gods assume when 
they travel. Also, ducks are associated with shamanism, 
perhaps because, as wide-ranging water birds, they ap- 
pear to inhabit several planes at once. 

Of even greater interest isa Monte Alban effigy snuff 
ing pipe of grey clay, representing a deer resting on its 
stomach, with legs drawn up and head turned to the 
right (Plate VIII). Cloven hooves leave no doubt about 
the zoological identification. What makes this piece 
especially fascinating is that it holds an unmistakable 
peyote cactus in its mouth (I am greatly indebted to 
Miss Julie Jones of the Museum of Primitive Art, 
who recognized the significance of the deer-peyote asso- 
ciation here in relation to the Huichol conceptualization 
of peyote as deer, and vice versa, and who, on that ac- 
count, drew my attention to the artifact). 

While anthromorphic pipes from Oaxaca and Nochi- 


[ 18 J 


Puiate VII 


Black-slipped, burnished terracotta efigy snuffer in the form of a 
duck-bodied man, from Monte Alban, Oaxaca, Late Formative, Monte 
Alban I[A-II (2), ca. 300-100 B.C. L. 33/, Anon. private collection. 
A number of similar snuffers combining head and animal characteris- 
tics, with the nose piece either in the tail or forming a horn on the 
head, are in the Museo del Arte Zapoteca, Mitla, Oaxaca. 


pala have the nosepiece in the form ofa horn on the head, 
it is the tail that forms the nosepiece in the deer efligy 
snuffer from Oaxaca. Such choices on the part of the 
prehistoric pipe makers cannot be considered to be arbi- 
trary. If one may venture some guesses, the horn atop 
the head as nosepiece may relate to the well known and 
widespread concept of horns—both single and double— 
as asymbol and even a source of shamanic or supernatu- 
ral power (I*urst 1965). Single horns on the forehead are 
a characteristic especially ofa certain class of Colima figu- 
rines, but they also occur elsewhere in Mesoamerica (e. g. 
at Tlatileo, Tlapacoya, Chalcatzingo, Nochipala, Monte 
Alban, ete.) and even in Peru, especially in Nazea art. 
As for the nosepiece of the deer effigy pipe, this might 
have to do with the concept of the deertail as magical 
power object in some North American shamantic prac- 
tices and beliefs. Among the Huichol, for example, the 
deertail is an important element in the shaman’s equip- 
ment, as it is in Papago shamanism. Likewise, it is 
hardly insignificant that the name of one of the principal 
Huichol supernaturals is Tamatst Mava Kwaat, Elder 
Brother Deer ‘Tail. 

The association of deer, divine inebriant and shaman 
which we perceive archaeologically in the Monte Alban 
snuffing pipe and ethnographically in Huichol and Cora 
religion, is itself an important culture-historical problem 
that remains to be seriously explored. Andean art dating 
to the fifth or sixth century A.1). suggests that there 
was something very like these Mesoamerican associations 
also in Peru. A common theme on Moche IV painted 
ceramics is a ritual deer hunt, in which the hunter is 
clearly not meant to be an ordinary man but a god, cul- 
ture hero, or great shaman (Plate IX). Moche painters 
consistently depict the deer in association with a shrub 
or tree which, though to some degree conventionalized, 


[ 20 | 


PuLaTE VIII 


Effigy snuffing pipe of burnished grey clay in the form of a deer 
holding a peyote cactus in his mouth, from Monte Alban, Oaxaca, 
Late Formative, Monte Alban I-II, ca. 300-100 B.C. The erect tail 
of the animal forms the nosepiece. Anon. private collection. L. 43’. 


[ 21 | 


is identifiable botanically as cfmadenanthera colubrina, 
with the long, bean-like seed pods characteristic of the 
hallucinogenically rich family Leguminosae. The seeds 
of this tree, called vilea or willka in the Andes, are made 
into a potent psychotomimetic snuff; they are also in- 
gested in a beverage and, in some highland Quechua 
villages, play an important role in the making of //ampu, 
a sacred substance used in cattle increase ceremonies and 
other rituals the origins of which lie far back in Andean 
prehistory (Billie Jean Isbell, personal communication). 

In any event, the deer is often a semi-divine celestial 
animal for American Indians, connected with Sun, Fire, 
sky beings, and shamans. Among the Warao of the 
Orinoco Delta, its flesh is still strictly taboo for shamans, 
suggesting at least a former sacred relationship (Johannes 
Wilbert, personal communication). Among the Huichol, 
it is the shaman’s spirit helper and companion: a pair of 
feathered ceremonial arrows that he wears on his head 
in certain ritual contexts symbolize deer antlers; the 
oblong basket of shamanic power objects (takwatsi) 1s 
identified with the divine Deer Person, MKauyumarie; 
certain deities are deer and vice versa; the deer is mount, 
guardian and guide on the shaman’s celestial quests and 
fights, especially on the peyote hunt; the ‘*Principal 
Deer’, Elder Brother Waiwatsari, is peyote, and vice 
versa, etc. 

Such concepts remind one at once of the role of the 
deer in Paleo- Asiatic or Siberian shamanism. In Siberia, 
too, the deer is the celestial mount that carries the sha- 
man to the Upperworld and its spirit rulers. In parts of 
Siberia, moreover, there is direct association between 
deer and the divine inebriant 
used by shamans to attain the ecstatic trance states in 
which they embark on their supernatural journeys—the 


in this case the reindeer 


Amanita musearia, or fly agaric mushroom, for which 


[22] 


PuatE [LX 


Supernatural deer hunting scene on a Mochica IV terracotta vessel, 
northern Peru, ca. A.D. 500. Drawn from the original by Alan 
Sawyer. Here, as in similar Mochica deer hunting scenes, the animal] 


is shown in association with a tree or shrub almost certainly to be 
identified by its characteristic seed pods as Anadenanthera colubrina, 
whose seeds were, and still are, ground into a potent psychotomimetic 
snuff, widely known as vi/ea or willka. In some Andean communities 
vilca seeds are also used in an intoxicating ceremonial drink; else- 
where, as in Chile, the pods may be burned and the smoke inhaled 
to achieve ritual intoxication. 


the reindeer is said to have an inordinate predeliction 
(Wasson 1972:204) and which some scholars regard as 
the Paleolithic or Mesolithic prototype for the Meso- 
american mushroom cults. 

It is difficult to escape the conclusion that the esteem, 
not to say veneration, with which some American In- 
dians regarded the deer represents a survival from an 
ancient, archaic, shamanistic substratum—a substratum 
that forms the underlying basis of American Indian 
ideology, including that of Mesoamerican civilization, 
and the ultimate roots of which he in the religion of 
Eurasian Paleolithic and Mesolithic hunting and gather- 
ing culture. ‘The curious association of deer as celestial 
mount on the shaman’s ecstatic Journeys and the sacred 
hallucinogens that are employed as aids in such mystical 
quests in parts of northern Asia as well as in America 
might well be a part of this very ancient belief system. 

In this connection, a new series of radiocarbon dates 
from rock shelter sites in Trans-Pecos Texas and north- 
ern Mexico is of special significance. These dates, for 
which IL am indebted to J. M. Adovisio of the University 
of Pittsburgh‘, confirm a time depth of over ten thousand 
years for the use of the potent hallucinogenic red seeds 
of the Sophora secundifiora shrub by Desert Culture 
hunters and food collectors as well as historic tribes in 
the same area. More than that, one important and well 
studied Texas site, known as Bonfire Shelter, yielded 
Sophora secundiflora from the lowest occupational stra- 
tum—Bone Bed II, witha C" age of 8440 to 8120 B.C., 
in direct association with Folsom and Plainview pro- 
jectile points and the bones of extinct bison. The same 


'A4 short paper on the topic by J. M. Adovisio and G. F. Fry was 
presented at the 71st Annual Meeting of the American Anthropol- 
ogical Association, Toronto, Canada, November 1972. A fuller treat- 


ment by the authors is in preparation. 


[24] 


seeds also occurred in the topmost occupational level, 
dated A.1). 420 to 1040, and in all of the intervening 
cultural deposits. A related rock shelter site in northern 
Mexico, Frightful Cave, similarly yielded Sophora se- 
cundifiora beans from its lowest level, dated at 7500 B.C., 
through all subsequent cultural deposits. Interestingly 
enough, here, as in many other Desert Culture rock 
shelter sites, Sophora secundiflora was invariably asso- 
ciated with Ungnadia speciosa, in contexts strongly sug- 
gesting, according to Adovisio, ritual use. 

It was Weston La Barre who suggested on several 
occasions (e.g. 1972: 270-278) that the origins of the 
American Indian hallucinogenic complex had to be 
sought ultimately in ecstatic, vision-seeking Paleoasiatic 
shamanism, the fundamental religion of the pre- 
agricultural Paleolithic and Mesolithic hunting peoples 
of Siberia who presumably constituted the ancestral pool 
from which flowed the Late Pleistocene migrations into 
North America. La Barre’s contention, which came to 
be increasingly shared by some of his colleagues in the 
study of aboriginal American religion and the botany and 
anthropology of hallucinogens, thus appears to be con- 
firmed: the historic shamanistic ‘‘red bean”’ cult of the 
Southern Plains is at least as old as the big-game hunt- 
ing phase of the terminal Pleistocene and thus appears 
to reach back toward a time when the peopling of North 
America across the Bering land bridge might still have 
been in progress. If knowledge of ritual or divine in- 
ebriation with plant hallucinogens was part and parcel 
of the intellectual baggage of these early migrants, it 
implies at least equal antiquity, if not a much greater 
one, for such practices in Eurasia. 

Just when or where in prehistoric antiquity the tech- 
nology and chemistry of snufling might have arisen is 
unknown. The concentration of these variants on the 


[ 25 | 


common theme of divine inebriation especially in the 
Amazon suggest that area as its ultimate source. If it 
did indeed diffuse from the tropical lowlands, its age 
would have to be at least three and a half thousand years, 
since, as mentioned above, the earliest known snufhing 
implements date to the mid-second millenium B.C. 
The fact that in addition to its numerous other psycho- 
tomimetics prehispanic Mesoamerica can now be shown 
to have shared in a wider pan-.American complex involv- 
ing the use of snuff opens up a host of new possibilities 
for culture-historical and ethnobotanical research. Ob- 
viously, multidisciplinary study of what now appears to 
have been a Mesoamerican snufling complex of substan- 
tial distribution and duration is important in and of itself. 
At the same time, it might provide answers to questions 
of external relationships, especially with South America, 
at roughly that crucial moment in time when Neolithic- 
type farming communities became transformed, by some 
as yet little understood processes, into the first great 
Mesoamerican civilization—that of the Olmec. 


REFERENCES 


Adovisio, J.M. and G.F. Fry. 1972. “‘Prehistoric Psychotropic Drug 
Use in Northeastern Mexico and Trans-Pecos Texas.”’ 
pared for the 71st Annual Meeting of the American Anthropological] 
Association, Toronto, Canada, 1972. 


Paper pre- 


Furst, Peter T. 1965. “West Mexican Tomb Sculpture as Evidence 
for Shamanism in Prehispanic Mesoamerica.’” Anthropologica 15: 
29-60. Caracas. 


——, 1968. ‘The Olmec Were-Jaguar Motif in the Light of Ethno- 
graphic Reality.’’ In: Dumbarton Oaks Conference on the Olmec, 
Elizabeth P. Benson, ed., pp. 143-178. Washington, D.C.: Dum- 
barton Oaks Research Library and Collection. Trustees for Har- 
vard University. 


——, 1972. “Ritual Use of Hallucinogens in Mesoamerica: New 
Evidence for Snuffing from the Preclassic and Early Classic.’’ In: 
Religion en Mesoamerica, X11 Mesa Redonda, Sociedad Mexicana 
de Antropologia, pp. 61-68. 


Gay, Carlo T.E. 1972. NXochipala: The Beginning of Olmec Art. The 
Art Museum, Princeton. 


Hissink, Karin, and Albert Hahn. 1961. Die Tacana: Ergebnisse der 
Frobenius- Expedition nach Bolivien, 1952 bis 1954. Stuttgart: W. 
Kohlhammer. 


La Barre, Weston. 1972. ‘‘Hallucinogens and the Shamanic Origins 
of Religion.’’ In: Flesh of the Gods: The Ritual Use ef Hallucino- 
gens, Peter ‘I’. Furst, ed., pp. 261-278. New York: Praeger Pub- 
lishers, Inc. 


Reis Altschul, Siri von. 1972. The Genus Anadenanthera in Amerin- 
dian Cultures. Cambridge: Botanical Museum of Harvard Univer- 
sity. 

Schultes, Richard Evans. 1972a. “‘On Overview of Hallucinogens in 
the Western Hemisphere.’’ In: Flesh of the Gods: The Ritual Use 
of Hallucinogens, Peter 1. Furst, ed., pp. 3-54. New York: Prae- 
ger Publishers, Inc. 


——, 1972b. “Hex Guayusa from 500 A.D. to the Present. Etnolo- 
gisker Studier, No. 32, pp. 115-138. Stockholm. 


[27 ] 


r 2 + ~~ © Ory , wi * , 0 
Wassén, S. Henry. 1965. I'he use of some specific kinds of South 


American snuff and related paraphernalia.’’ Etnologisker Studier, 
No. 28, Stockholm. 


—-, 1967. “‘Anthropological Survey of the Use of South American 
Snuffs.”’ In: Ethnopharmacologic Search for Psychoactive Drugs, 
Daniel H. Efron, ed., pp. 233-289. Washington, D.C.: U.S. Pub- 
lic Health Service Publication No. 1645. 


1972, “The Anthropological Outlook for Amerindian Medi- 
cinal Plants.’? In: Plants in the Development af Modern Medicine, 
Tony Swain, ed., pp. 2-65. Cambridge, Harvard University Press. 


99 


Wasson, R. Gordon. 1972. *“What was the Soma of the Aryans?”’ 


In: Flesh of the Gods: The Ritual Use of Hallucinogens, Peter T. 
Furst, ed., pp. 201-213. New York: Praeger Publishers, Inc. 


otk 
oe 


_-<BOTANICAL MUSEUM LEAFLETS 
aa HARVARD UNIVERSITY 


Campringr, Massacnusertrs, Ocroper 25, 1974 VoL. 24, No. 2 


A STUDY OF SYSTEMATIC WOOD 
ANATOMY IN CANNABIS 
BY 
Loran C. ANDERSON! 


Cannabis has been associated with man since very early 
times (Ash, 1948), yet, surprisingly, little is known about 
its comparative wood anatomy. The reasons are due 
probably to the tendencies for (1) anatomists to select 
wood from trees and woody shrubs rather than from 
herbs for study and for (2) researchers often to disregard 
or slight plants associated with man, either as crops or 
weeds, in basic scientific enquiries. 

Tippo (1988) offered a few general comments on the 
wood of C. sativa L. in his extensive study on the anato- 
my of the Moraceae and its allies. Stem shape and leaf- 
trace number in transections were stressed by Nassonov 
(1940) in a report on geographical races of hemp. Hay- 
ward (1948) devoted a chapter in his textbook to C. 
sattva. The general morphology of that species was 
given, but details of seedling anatomy and floral struc- 
ture were emphasized; wood anatomy was scarcely men- 
tioned. Metcalfe and Chalk (1950) summarized anatomi- 
‘al data on Cannabaceae to that date. Shimomura e¢ al. 
(1967) emphasized trichomes in their study of leaf and 
bract anatomy in Cannabis; they found differences be- 
tween C. sativa and C. indica. 

' Department of Biological Sciences, Florida State University, Tal- 
lahassee, Florida. 


FCONOMIC BOTAI? 
OF CAKUS & 


CWT DA PTT OAT 


Rh het nd hae Ue 


Some features of Cannabis anatomy are relatively well 
known, suchas the economically important phloem (bast) 
fibres. These aspects have been reviewed by Hayward, 
1948: Metcalfe and Chalk, 1950. Considerable attention 
has also been given to cystolithic hairs (Pireyre, 1961) 
and laticifers in Cannabis. 

With the recent attention devoted to taxonomic prob- 
lems in Cannabis (Schultes et al., 1974: Stearn, 1974), 
I am pleased to present this introductory account on 
comparative wood anatomy. It includes apparently the 
first technical description of wood identified with vouch- 
ered material as C. ¢ndica Lam. 

Merruops AND Matrerias 

All materials were collected fresh and preserved in 
formalin-propriono-alcohol (FPA). Woods were sec- 
tioned on a sliding microtome at 20 h. Some sections of 
cach sample were stained in safranin © and counter- 
stained with fast green FCF and orange G: others were 
stained only with safranin. ‘Tissues were mounted in 
Permount. 

Xylem features were microscopically measured with a 
‘alibrated ocular micrometer: a minimum of 50 measure- 
ments were made for each feature reported in ‘Table 1. 
Polarizing filters aided study of cell wall structure and 
crystals. Statistical analyses were made on a Wang 600 
computer with the assistance of Dr. M. P. Johnson. 

The material of C. indica came from a wild population 
at Pashimool, west of Kandahar, Afghanistan, R. 1. 
Schultes 26505 (Keon. Herb. Oakes Ames); that of C. 
sativa came trom a naturalized population in Pottawato- 
mie County, Kansas, United States, 1. C. Anderson 
3668 (Fla. State Univ.). 

Resuirs 
Details of wood anatomy are illustrated in Figs. 1-6, 


30 | 


The woods of C. indica and C. sativa differ significantly 
in each feature listed in Table 1. 

Vessels in C. indica tend to occur in radial chains; 
whereas those of C. sativa usually occur singly (as illus- 
trated in Hayward, 1948). That difference in distribution 
can be seen in Figs. 1-2. Vessel members are angular 
to round in transection. ‘They have simple perforation 
plates, and the end walls are slightly oblique. Pits are 
alternate with elliptic borders. Pit apertures are elon- 
gate; they are 6-9 » long in C. indica and 4-8 p» in C. 
sativa. 

Vessel members and wood fibres differ between the 
two samples in average width, length and cell wall thick- 
ness (Table 1). In C. indica, both cell types are wider, 
have thicker walls, but are shorter in length compared 
to those of C. sativa. 

Fibres in the secondary xylem must not be confused 
with the hemp fibres of commerce, which are phloem or 
bast fibres. Wood fibres of C. indica are typical, lignified 
libriform fibres. Fibres in C. sativa differ in two respects. 
They are dimorphic, with successive tangential bands of 


Taste 1. Averaged measurements on wood anatomy in Cannabis. 


Feature C. indica C. sativa Significance 
level@ 
vessel number per group 3.05 1.39 ” 
vessel member width, / 68.52 62.16 * 
vessel member wall thickness, 4 3.50 2.30 = 
vessel member length, 209.71 244,54 at 
fibre width, /@ 18.41 14,28 eK 
fibre wall thickness, - 3.44 0.68 = 
fibre length, » 281.10 443.47 sii 
ray width (cell number) 2.23 1.63 a 
ray height, mm 0.87 0.68 * 


* Analysis of variance (F test): * = p < .05, ** = p < .001 


[ 31 ] 


EXPLANATION OF PLATE X 


Figs. 1-6. Cannabis wood sections. Figs. 1,3, 5 are C. indica, and 
2,4,6 are C, sativa. Fig. 1, transection showing radial chains of ves- 
sels, libriform fibres, and procumbent ray parenchyma. Fig. 2, tran- 
section showing tendency for solitary vessels, fibre dimorphism, and 
ray parenchyma shorter radially (erect). Fig. 3, transection showing 
thick walls of vessel members and libriform fibres ; note cuboidal crys- 
tals in ray cells (arrows). Fig. 4, transection showing relatively thin 
cell walls; note shrunken secondary walls of gelatinous fibres. Fig. 5, 
tangential section showing wood rays with numerous crystals; photo- 
graphed with partially polarized light. Fig. 6, tangential section 
showing relatively narrower wood rays with erect cells; crystals ab- 


sent. Figs. 1-2, 5-6, < 71. Figs. 3-4, < 308. 


[ 82 | 


PLATE X 


C. sativa 


indica 


C. 


* 
‘ 
s 
* 


e be 


2 


ob 
5 


See re © 


t 
‘ 


} 


* 


cnn nen EE 
oem 


[vceeteniessnsies ceeessemeeenttaiadidiiteensaiensanediiineeneenee 


onal Le aed 


thick-walled fibres alternating with bands of thin-walled 
fibres. They have irregularly shrunken secondary walls 
(more pronounced in the thick-walled fibres) and are 
termed gelatinous (Fig. 4). Their staining reaction (note 
lighter tones in Figs. 2, 4, 6) and absence of birefringence 
under polarized light are similar to that of gelatinous 
fibres in other species that I have studied (Anderson, 
1963, 1972). 

Axial parenchyma is paratracheal. It is very scanty in 
C. indica and scanty to vasicentric in C. sativa. 

Wood rays are classed as heterogeneous I: ¢.e., both 
multiseriates and uniseriates occur, and they are com- 
posed of procumbent and erect ray cells. Those of C. 
indica are predominantly square to procumbent ; whereas 
ray cells in C. sativa are mostly erect with very few square 
or procumbent ones. The differences in cell shape are sug- 
gested in Figs. 5-6, but they are best viewed in radial 
sections. A qualitative difference in wood rays is the 
presence of numerous cuboidal or prismatic crystals of 
‘calcium oxalate in C. indica. They can be seen in all sec- 
tions under normal light but are more obvious with par- 
tial polarization of light (Fig. 5). No crystals were found 
in C. sativa ray cells (although both species have druses 
in their phloem and ground tissues). 


Discussion 


Many American botanists have thought Cannabis to 
be monotypic, possibly because only hemp, C. sativa, 
has been cultivated in this country. Most taxonomists 
who have studied the genus closely, however, recognize 
three species: C. indica Lam., C. ruderalis Janisch., and 
C. sativa L. (see Schultes et a/., 1974, for a review of the 
taxonomic history of the genus). 

Data from wood anatomy have not hitherto been uti- 
lized in the taxonomy of Cannabis. Such data might 


[ 34 | 


help resolve the question of species recognition in the 
genus. Nassonov’s study (1940) is of little use, as he 
mentioned no binomials. He primarily studied variation 
in crop plants (all C. sativa’), where he identified three 
basic types of stem structure. He did note that wild and 
cultivated forms of hemp could not be distinguished 
clearly on the basis of anatomy of stem and bast fibres. 

Wood features of C. indica and C. sativa listed in 
‘Table 1 are those commonly measured in comparative 
studies. They are all significantly different between the 
species with four at the 5% level and five at the 0.1% 
level! Additional differences in the axial and radial paren- 
chyma systems are noted in the text. Woods of the 
two species are qualitatively distinct for libriform fibres 
versus gelatinous fibres and for presence of crystals in 
wood rays. Many examples of the taxonomic signifi- 
‘ance of crystals in woods have been noted (Bailey, 1961: 
Chattaway, 1955-56). 

Although only one sample of each species is discussed 
here, the magnitude of differences between the two is 
impressive in a system as conservative as wood. In his 
exhaustive review on many aspects of wood science, Jane 
(1963) stated the following regarding taxonomic wood 
anatomy: 


Wood structure is probably more conservative than floral struc- 
ture, and specific differences, as determined by floral characters, 
are often not reflected in the secondary xylem. Indeed, it may 
be said that in general the distinguishing features of wood are 


at generic, rather than specific, level. 


Certainly, the plants used in this study are of the same 
genus, but it is my opinion that they represent different 
species. 

examination of woods from three additional collec- 
tions of North American C. sativa shows they are also 
distinct from the C. 7ndica wood sample. All vary from 


C. indica in the features listed in Table 1, with the ex- 
ception of vessel member width. All three samples have 
gelatinous fibres. Crystals are absent in wood rays in 
two: only a few were found in the rays of the third 
sample. Complete data on these samples will be pre- 
sented in the future as part of an expanded study on the 
wood anatomy of Cannabis. 


LITERATURE CITED 

Anderson, L.C. 1963. Studies on Petradoria (Compositae): anatomy, 
cytology, taxonomy. Trans. Kans. Acad. Sci. 66: 632-684. 

—, 1972. Studies on Bigelowia (Asteraceae). I]. Xylary compari- 
sons, woodiness, and paedomorphosis. J. Arnold Arb. 53: 499- 
Olt, 

Ash, A.L. 1948. Hemp —production and utilization. Econ. Bot. 2: 
158-169. 

Bailey, I.W. 1961. Comparative anatomy of the leaf-bearing Cacta- 
ceae, III. Form and distribution of crystals in Pereskia, Pereskiop- 
sis and Quiabentia. 

Chattaway, M.M. 195 
Trop. Woods 102: 5 

Hayward, H.E. 1948. 
Co., New York. 

Jane, F.W. 1968. Botanical aspects of wood science. /n: W.B. Tur- 
rill, ed., Vistas in Botany, Vol. I]. Pergamon Press, Oxford. 

Metcalfe, C.R., and L. Chalk. 1950. Anatomy of Dicotyledons, Vol. 
II. Oxford Univ. Press, London. 

Nassonov, V.A. 1940, “Anatomical characteristics of geographical 
races of hemp’’ [in Russian]. Vestnik Sotsialisticheskogo Rasten- 
ievostva, Institut Rastenievostva, Moscow, 4: 107-120. 

Piereyre, N. 1961. Contribution a l°étude morphologique, histologi- 


56. Crystals in woody tissues. I and II. 
74, 104: 100-124. 
he Structure of Economic Plants. Macmillan 


5 
9) 
T 


que et physiologique des cystoliths. Rev. Cytol. et Biol. Vég. 23: 
93-320. 

Schultes, R. K., W.M. Klein, T. Plowman, and T. EK, Lockwood. 
1974. Cannabis: an example of taxonomic neglect. Harvard Univ. 
Bot. Mus. Leafl. 23: 337-367. 

Shimomura, H., M. Shigehiro, KE. Kuriyama, and M. Fujita. 1967. 
Studies on Cannabis. 1. Microscopical characters of their internal 
morphology and spodogram. J. Jap. Pharm. Soc. 87: 13834-1341. 

Stearn, W.T. 1974. Typification of Cannabis sativa L. Harvard Univ. 
Bot. Mus. Leafl. 23: 325-336. 

Tippo, O. 1938. Comparative anatomy of the Moraceae and their 


presumed allies. Bot. Gaz. 100: 1-99. 


TWO NEW BRAZILIAN SPECIES 
OF BRUNFELSIA 
BY 
Timotuy C. PLowMaAaNn 


In anticipation of my forthcoming revision of the 
South American species of Brunfelsia (Solanaceae) based 
on a doctoral dissertation at Harvard University (Plow- 
man, 1973), [ herewith offer the description of two previ- 
ously unrecognized species. 

I would like to express my thanks to the curators of the 
several herbaria who have graciously loaned specimens 
cited in this paper. Abbreviations of herbaria are taken 
from Index Herbariorum (I.anjouw and Stafleu, 1964). 
Special thanks are due also to Dr. Leslie Garay for 
checking the Latin descriptions and to Lynda Bates for 
preparing the line drawings which accompany the text. 

Research reported in this paper was supported by the 
National Institutes of Health Training Grant (T TO] 
GM 00036-13) and by the National Science Foundation 
Evolutionary Biology Training Grant (GB 7346, Reed 
Rollins, Principal Investigator, Harvard University). 


Brunfelsia Martiana Plowman sp. nov. 

Frutex. Folia subsessilia, oblonga, plerumque elliptico- 
oblonga vel oblongo-lanceolata, apice acuminata, basi late 
cuneata vel obtusa, utrinque glabra, nervis lateralibus 
8-12, petiolo brevissimo. Inflorescentia terminalis vel 


[ 37 ] 


L 


axillaris annotinis ramulis, pauciflora, glabra. Flores 1-7, 
albidi vel albo-virescentes. Pedicellus erectus, gracilis, 
tlaber. Calyx tubulosus vel tubuloso-campanulatus, 


( 
Sb 
( 


| 
glaber, dentibus subaequalibus, triangulari-ovatis vel 
ovato-lanceolatis, apice acutis vel acuminatis. Corollae 
tubus quam calyx duplo longior, rectus, apice parum in- 
flatus, glaber, limbo patent, lobis subaequalibus, oblongo- 
obovatis, apice rotundatis vel subtruncatis, marginibus 
lateralibus abrupte reflexis.. Stamina inclusa in tubi parte 
superiori. Ovarium oblongo-ovoideum, Fructus ignotus. 

A Brunftelsia guianense foliis oblongis, longioribus, 
nervis lateralis pluribus (8-12), calyce tubuloso-campan- 
ulato et corollae tubo quam calyce duplo longiori et a 
B. amazonica foliis majoribus et pedicellis multo breviori- 
bus differt. 

Tyre: Brazil: Amazonas: ‘‘Provinciae Rio Negro, habitat in sylvis 
ad flum. Japura, prope Sao Joao do Principe,’ Dec. 1819, Martius 
[3247] (holotype, M; isotypes, M). 

Shrub to 1m. tall. Branches few, naked below, some- 
what knobby at nodes, glabrous. Bark cracked longitu- 
dinally, dark reddish brown, shiny, furnished with len- 
ticels. Leaves scattered along branchlets, subsessile, 
mostly oblong, sometimes elliptic-oblong or oblong- 
obovate, acuminate at apex, broadly cuneate to blunt 
at base, blade 10-25 cm. long, 4-8 cm. wide, glabrous on 
both surfaces, firmly membranaceous to subcoriaceous, 
upper surface dark green, dull or shiny, lower surface 
paler green, sometimes shiny, midrib dark reddish brown, 
lateral nerves 8-12 pairs, spreading, straight: petiole 
short, 1-4 (12) mm. long, glabrous, becoming cracked, 
rugose. Inflorescence terminal or axillary in the upper 
leaf axils, glabrous, axis 2-10 (15) mm. long. Flowers 
1-7 per inflorescence, sometimes with short peduncle 
which articulates with pedicel, greenish white to white. 
Bracts 1-8 per flower, linear-lanceolate, concave, 1-10 


PLATE XI 


| Sic spigec i p 


Per 9? she 


TYPE 


HOLOTL EE festeton of Brvntzfee ‘ Soinne 
Brusfelela gent jan 


| Me Meirima Sore sey A | 2 a8 


Photograph of holotype of Brunfelsta Martiana Plowman 


K.F.P. von Martius [3247]. 


mm. long, ciliolate at margin, caducous. Pedicel short, 
slender, 8-6 (10) mm. long, glabrous. Calyx tubular to 
tubular-campanulate, 8-12 mm. long, 8-8 mm. in di- 
ameter, glabrous, rarely striately veined, teeth subequal, 
triangular-ovate to ovate-lanceolate, 2-6 mm. long, acute 
to acuminate at apex. Corolla tube twice as long as 
calyx, straight, cylindric, inflated at apex, 20-24 mm. 
long, 1-8 mm. in diameter, glabrous; limb spreading, 
somewhat undulate, 15-22 mm. across, lobes subequal, 
oblong-obovate, abruptly reflexed at lateral margins, 
rounded-truncate at apex, 5-10 mm. long. Stamens in- 
cluded within upper part of tube; filaments subligulate, 
longer anterior pair 4 mm. long, shorter posterior pair 
3 mm. long; anthers orbicular-renitorm, slightly un- 
equal, the upper pair somewhat smaller, about 1 mm. in 
diameter. Ovary oblong-ovoid, 2 mm. long: style fila- 
mentous, a little broader and curved at apex, about 15 
mm. long: stigma briefly bifid, upper lobe slightly 
larger, 1 mm. long. Fruit and seed unknown. 


Disrrrpurion. Brazil (Amazonas, Para, Bahia); Guyana. 

AppITIONAL specIMENS. Brazit: Para: Faro, 22 Jan. 1910, Ducke 
s.n. (MG): Bella Vista, Rio Tapajoz, matta das immediacoes do 
Campina do Perdido, 12 Feb. 1917, Ducke s.n. (MG): Regiao do 
Jutahy de Almeirim. Palhal, 16 Apr. 1923, Ducke s.n. (RB); Gurupa, 
25 Feb. 1923, Ducke s.n. (RB); Parad, May 1819, Martius 3300 (M). 
AMAZONAS: Manaus and vicinity, road Manaus-Caracarai, km. 22, 
21 Mar. 1967, Prance et al., 4704 (WIS), Rio Negro, Cachoeira Baixa 
de ‘T'arumao, 11-14 Apr. 1973, Schultes & Rodrigues 26132-A (ECON), 
26188 A(ECON). Banta: Distrito Ileus, Ferradas, Dec. 1818, Mar- 
tius s.n. (M): Feira de Santana, Apr. 1850, collector unknown (G). 

Guyana: Essequibo-Demarara River, Madray-Bubu Trail, Topy 
Trysil Forest, 8 Feb. 1944, Forest Department 4422 (K). 


Brunfelsia Martiana is named for K.F.P. von Martius 
who first collected the plant in the year 1818. This new 
specific concept has proven difficult to define, not through 
its lack of distinctive characters, but due to the scarcity 
of specimens which are scattered in Brazilian and Euro- 
pean herbaria. 

[ 40 | 


Piate XII 


BRUNFELSIA = martiana Plowman 


Brunfelsia Martiana Plowman. 1, flowering branch, one third natural size. 2, 
corolla limb from beneath, approximately natural size. 3, calyx showing attach- 
ment to stem and bracts, approximately natural size. 4, excised corolla tube 
showing stamens and pistil, one and one-half times natural size. 


This interesting shrub grows in the understory of low- 
land tropical forests from Guyana throughout the lower 
Amazon basin and with disjunct populations in the 
coastal forests of the State of Bahia. The species has 
been collected only twice in the last fifty years, and no 
collections of it are known from Bahia since 1850. 

Brunfelsia Martiana is closely related to B. amazonica 
Morton and B. guianensis Benth., which also have 
rather small, greenish white flowers and occur sym- 
patrically with 2B. Martiana in the middle Amazon. 
B. Martiana differs from 2B. amazonica in having larger 
leaves (10-25 em. vs. 6-12 em.) with more lateral nerves 
(8-12 pairs vs. 5-8 pairs) and much shorter pedicels 
(8-6 mm. vs. 13-20 mm.). It differs from B. euianen- 
sis in having larger (10-25 cm. vs. 6-15 em.) oblong 
leaves with more lateral nerves (8-12 pairs vs. 4—5 pairs), 
a calyx which is usually narrow-tubular, not ovoid- 
‘ampanulate, and a corolla tube which is rarely more 
than twice as long as the calyx. In B. euranensis, the 
tube is frequently 2) to 8 times as long as the calyx. 


Brunfelsia pilosa Plowman sp. nov. 

Krutex. Ramuli graciles, pilosi vel villosi. Folia breve 
petiolata, anguste elliptica, oblongo-lanceolata vel obo- 
rata, apice acuminata, basi cuneata, utrinque ad costam 
pilosa, nervis lateralibus 6-9. Inflorescentia terminalis, 
sessilis. Flores 1-8, speciosi. Pedicellus brevissimus, 
apicem versus incrassatus, pilosus. Calyx tubuloso- 
ventricosus, in exsiccatione campanulatus, sparse vel 
dense pilosus, raro glaber, membranaceus, dentibus sub- 
aequalibus, lanceolatis, acuminatis; calyx in fructu per- 
sistens, quam capsula fere aequans. Corollae tubus quam 
‘calyx duplo longior, glaber, limbo patenti, lobis sub- 
aequalibus, rotundatis, lobo superiori posterior! parum 
majori. Stamina et stigma in tubi parte superiori in- 


[42] 


Piare XIII 


Photograph of holotype of Brunfelsia pilosa Plowman, L.B. Smith 
& Pe. R. Reitz 12777. 


» 


= 
cs 
. 
Ne 
ee 


clusa. Ovarium conico-ovoideum, Capsula ovoidea vel 
subglobosa, apice apiculata, laevis, atroviridis. Semina 
ellipsoides, reticulato-foveata. 

A Bruntelsia cuneifolia calyce tereti pubescentia pilosa 
et corollae tubo longiori, a B. uniflora calycis dentibus 
longioribus, pubescentia pilosa et corollae tubo longiori, 
et a B. australe foliis apice acuminatis, pubescentibus 
angustioribus differt. 

Tyrer. Brazit: Santa Catarina: Municipio Sao Miguel d’Oeste, 
forest above Rio Reperi-guacti, Peperi, ca. 26°32’ S., 53°44’ W., 
300-400 m., 21 Oct. 1964, LL.B. Smith & Pe. R. Reits 12777 (holo- 
type, GH; isotypes, MO, R, UC). 


Shrub 0.5—-2 m. tall, diffusely branched from near 
base. Branches spreading and arching, terete. Bark 
rough, longitudinally cracked, shedding in thin, char- 
taceous flakes, yellowish brown. Branchlets slender, 
pilose to villous, greyish green, dark purple when young. 
Leaves appearing two-ranked, scattered on branchlets, 
narrowly elliptic, oblong-lanceolate or obovate, acumi- 
nate at apex, cuneate at base, blade 8-7.5 cm. long, 
1.8-8 em. wide, pilose on both surfaces, primarily at 
midrib, firmly membranaceous, dull, dark green above, 
pale green beneath, the young leaves dark purple, lateral 
nerves 6-9 pairs, straight, often prominulous above; 
petiole short, 1-4 mm. long, pilose to villous. Inflores- 
cence terminal, sessile, usually with one flower, rarely 
2-3. Flowers showy, deep violet fading to pure white 
with age, odorless. Bracts 1-8, linear-lanceolate, 1-8 
mm. long, pilose to villous, caducous. Pedicel short, 
stout, 1-8 mm. long, pilose; in fruit becoming corky, 
rugose-verrucose toward apex. Calyx tubular-ventricose, 
appearing campanulate in pressed specimens, terete in 
cross-section, 12-19 mim. long, sparsely to densely pilose 
with long weak hairs, rarely glabrous, membranaceous, 
purplish, drying reddish brown, teeth subequal, lanceo- 


[ 44] 


PLATE XIV 


BRUNFELSIA pilosa Plowman 


Brunfelsia pilosa Plowman, 1, flowering and fruiting branch, one- 
half natural size. 2, corolla limb from above, approximately natural 
size. Drawn from living plant in cultivation at Miami, Florida, Plow- 
man 2963 (ECON). 


(45 | 


late, 4-10 mm. long, acuminate; calyx in fruit persist- 
ent, partially enclosing capsule, becoming subcoriaceous, 
light green, the sinuses becoming more deeply cut. Co- 
rolla tube twice as long as calyx, 25-82 mm. long, 1.5-8 
mm. in diameter, glabrous; limb spreading, 30-47 mm. 
in diameter, thickening at mouth prominent, round and 
white, lobes subequal, the uppermost posterior lobe 
slightly larger, broadly rounded, 10-15 mm. long. Sta- 
mens included in uppermost part of corolla tube; fila- 
ments slender, upper pair 38-4 mm. long, lower pair 38-5 
mm. long, glabrous; anthers orbicular-reniform, 1—1.5 
mm. in diameter. Ovary conical-ovoid, 2—2.5 mm. long, 
1 mm. in diameter; style slender, 22-26 mm. long: stig- 
ma briefly bifid, in the form of a forceps, 1 mm. long. 
Capsule ovoid to subglobose, apiculate at apex, 12 mm. 
long, 10 mm. in diameter, smooth, dark green, shiny, 
pericarp thin-walled, dry at maturity, tardily dehiscent. 
Seeds ca. 10, ellipsoid, 5-6 mm. Jong, 2.5-4 mm. in 
diameter, reticulate-pitted. 


Distrisution. Brazit: (Sao Paulo, Parani, Santa Catarina, Rio 
Grande do Sul); Paraguay ; Argentina (Misiones). 

AbbDITIONAL specIMENS. Brazit: Sao Paulo: Itapetininga, 17 Nov. 
1887, Léferen 389 (C, R, SP, US); Ypiranga, Nov. 1910, Liiderwaldt 
2717 (RB, SP); Campinas, Jundiahy, Mar. 1900, Campos Novaes 216 
(US); Villa de Serra Branca, 1 Nov. 1897, Puttemans 4328 (SP); 
Carandirti, Dec. 1912, Tamandaré 244 (RB); Mandaquil, 23 Nov. 
1906, Usteri s.n. (G), Parana: Curitibi, Bairro Sao Niolau, 18 Nov. 
1966, Capriliont 1659 (US); Itaperussti, 17 Nov. 1908, Dusén 7077 
(GH, S); Jaguariahyva, 25 Oct. 1910, Dusén 10443 (GH, MICH, 
NY, S): Tamandaré, 4 Oct. 1914, Fénssen 1054a (F, GH, NY, S$); 
Sao Matheus, 27 Feb. 1929, Gurgel s.n. (RB): Municipio Rio Branco 
do Sul, Sao Vicente, 27 Oct. 1967, Hatschbach 17610 (C, UC); Ponta 
Grossa, 2 Nov. 1928, Hoehne 23309 (SP, US); Foz do Iguacti, Parque 
Nacional de Iguacti, 8 Oct. 1946, J.G. Kuhlmann s.n. (RB) :Ypiranga, 
15 Sept. 1934, Reiss 99 (GH, NY). Santa Catarina: Municipio 
Cacador, Fazenda dos Carneiros, 1100 m., 7 Dec. 1962, Aletn 3518 
(US); Municipio Campos Novos, Palmares, 950 m., 28 Oct. 19683, 
Klein 4102 (US); Bituruna, Fazenda Etienne, 11 Feb. 1948, Mello 
Filho 793 (R); Nova Teutonia, 25 Oct. 1943, Plaumann 164 (RB); 


[ 46 ] 


Municipio Ararauja, Rodeio da Areia, 22 Nov. 1943, Reits C-171/ 
(RB); Santa Cecilia, 1100 m., Reitz & Klein 14136 (NY): Municipio 
Sao Miguel d’Oeste, Canela Gaucha, 8 km. northwest of Sao Miguel 
d’Oeste, ca. 26°40! S., 53°34’ W., 700-750 m. , Smith & Reitz 12757 
(NY, R, US); Manic pio: Abelardo Luz, north bank of Rio Chapeco 
at Abelardo Luz, ca. 26°35 S.; 52°20/ W., 900-1000 m., 28 Oct. 
1964, Smith & Reitz 12870 (C, F, ae. R); Tubarao, Nov. 1889, Ule 
1521 (P). Rio Grande do Sul: Sao Leopoldo, 10 Sept. 1946, Henz 
27246 (MO, NY), 20 m., 17 Sept. 1946, Henz 35305 (S): Municipio 
Rio Pardo, Fazenda Horticola, 70 m., Oct. 1923, Jiirgens 19 (B); 
Colonia Santo Angelo, 4 Feb. 1898, Lindman A-597-b (S), Sept. 
1900, Schwarzer 50(L,S); Silveira Martins, 20 Mar. 189 3, Lindmann 
A-597-c (S); Rio dos Sinos, 8 Nov. 1949, Rambo 44295 (BR, L); 
Bitterberg prope Montenegro, 13 Nov. 1950, Rambo poy (GH 
P, W). 

Paracuay: In regione fluminis Alto Parana, Yaguarazapa, 1909- 
1910, Frebrig 5460 (G, GH, US). 

ARGENTINA: Misiones: Departamento de San Javier, Acaragua, 
220 m., 30 Sept. 1947, Bertoni 2974 (B, W); Departamento de Can- 
delaria; Bompland, 4 Oct. 1909, Jorgenson s.n. (BAB), Nov. 1910, 
Jorgenson s.n. (BAB); Santa Ana, Aug. 1901, Llamas 1530 (BAB, 
G). Departamento de Iguazt, Cataratas de Iguazu, 13 May 1969, 
Plowman 2735 (GH); Delicia, camino a El Dorado, 4 Nov. 1949, 
Schwindt 2270 (C, LD, NY); Puerto Aguirre, Rio Iguazti, 19 Sept. 
1922, IL.N.T.A. 44365 (B). 


Brunfelsia pilosa is a distinct species which has been 
consistently confused with several other plants of south- 
ern Brazil, especially B. uniflora (Pohl) Benth. and B. 
australis Benth. An early collection of B. pilosa (Sellow 
1573) with atypically glabrous leaves and calyces was 
cited by Bentham as a syntype of his concept of B. 
australis (Bentham, 1846). 

A small shrub with showy flowers, Brunfelsia pilosa 
occurs primarily in the understory of primary and second- 
ary woods, as well as in thickets and along water courses. 
Flowering takes place from October to December with 
fruits appearing from February to May. It has a fairly 
wide altitudinal range, occurring from near sea level to 
1100 m. ‘The species ranges from the Brazilian State of 
Sao Paulo south to Rio Grande do Sul and extreme 


[ 47 ] 


northeastern Argentina (Misiones). In the central part 
of its range, where it occurs most commonly, this species 
seems to be associated with the formerly extensive 4 rau- 
caria forests of the south Brazilian planalto. 

Owing to its attractive flowers, this plant is cultivated 
in tropical gardens, although not so extensively as Brun- 
felsia australis. Recently, B. pilosa has appeared in the 
Florida nursery trade and iS increasing in importance as 
an ornamental. 

Brunfelsia pilosa may be distinguished from related 
species by its very short (1-8 mm. long), stout pedicels 
and an inflated tubular calyx with deeply cut teeth 
(4-10 mm. long). The twigs, leaves and calyx charac- 
teristically contain few to many long, weak trichomes. 
The flowers are large and showy with a corolla limb 
nearly 5 cm. in diameter. 


LITERATURE CITED 
Bentham, G. m A. DeCandolle, 1846. Prodromus Systematis Natu- 


ralis Regni Vegetabilis. 10: 200. 


Lanjouw, J. and F.A. Stafleu, 1964. Index Herbariorum. 5th Fd. 
Regnum Vegetabile Vol. 31. International Bureau for Plant Tax- 
onomy and Nomenclature, Utrecht. 


Plowman, T. 1978. The South American Species of Brunfelsia (So- 


lanaceae). Doctoral Dissertation, Harvard University, Cambridge, 
Massachusetts. 


[ 48 | 


BOTANICAL MUSEUM LEAFLETS 


HARVARD UNIVERSITY 


CaMBRIDGE, MAssacnuseTTs, JANUARY 10, 1975 Vou. 24, No. 3 


ARCHAEOLOGICAL MAIZE FROM 
NORTHERN CHILE 
BY 
Paunt C. Mancetsporr AND Gorpon C. PoLLarp* 


INTRODUCTION 


Now that the modern races of maize of the countries 
of this hemisphere have been classified and described in 
a series of eleven publications issued by the National 
Academy of Sciences- National Research Council, it has 
become a common procedure for botanists analyzing 
collections of prehistoric remains of corn of a particular 
country to relate the ancient specimens, so far as is possi- 
ble, to the living races of that country. The modern 
maize of Chile has been described by Timothy ef al. 
(1961), who recognized nineteen more or less distinct 
races. 

As part of a study of prehispanic cultural development 
in the Atacama Desert of northern Chile, the junior 
author of this report obtained several collections of maize 
cobs from archaeological sites found near the town of 
Chiu Chiu, which lies beside the middle section of the 
Loa River. The vicinity of Chiu Chiu, at an elevation 
of 2500 meters, is a small oasis along the river which 
supports a narrow band of vegetation in the extreme 


* Associate Professor of Anthropology, State University of New 
York College at Plattsburgh. 


[ 49 ] 


desert environment. The region around the town has a 
long and almost unbroken archaeological sequence begin- 
ning at least 10,000 B.C. and lasting until post-Conquest 
times. A summary of the cultural sequence since the in- 
troduction of ceramic technology (ca. 500 B.C.) is found 
in Pollard (1971). The appearance of maize cultivation 
in the archaeological record (ca. A.D). 100) is associated 
with artificial irrigation technology. 


MalIze FROM THE Earnikst SITE 


The earliest site found with associated maize is desig- 
nated RAnL 100. Although no prehispanic structures 
are preserved, the site is believed to be the earliest agri- 
cultural village settlement along the middle Loa River. 
Sections of the site include a small intact refuse mound, 
a looted cemetery, ancient agricultural terraces and irri- 
gation canals, and ruins of a historic-period house. -A 
1.0 & 2.5 m. cut was excavated in the refuse mound, 
revealing five well separated, but culturally homogeneous 
layers, the deepest of which (layer 5) was as much as 55 
em. below ground surface. Each layer yielded an abun- 
dance of sherds, various lithic artifacts, and = sizeable 
quantities of Lama sp. bones. Layers 2-5 each con- 
tained varying amounts of Lama sp. wool and mesquite 
seeds (Prosopis chilensis). Layers 8 and 5 also contained 
several gourd fragments (Lagenaria sp.). Twenty maize 
cobs and several fragments were recovered from layers 
2—5. Radiocarbon analysis of asample of ama sp. wool 
and skin from layer 5 resulted ina date of A.D. 105 + 
105 (GN-1644). At present, this is the earliest date for 
maize agriculture in northern Chile. 

Of the nineteen races of maize described by ‘Timothy 
et al. (1961), we recognize three in the collection trom 
site 100. Descriptions of these follow : 


[ 50 | 


Capio Chico Chileno 


Twenty of the oldest cobs are considered to be related 
to this modern race; their characteristics are set forth in 
Table 1. All of the cobs represented in this table came 
from the test excavation described above. Layer pro- 
venience is represented by the last number in the code, 
except that layer 2 is represented by two code numbers, 
2 and 7. There is no clear relationship between the depth 
of the layer and the characteristics of the cobs. Within 
each layer the cobs are arranged in order of their lengths, 
for identification purposes. 

We call particular attention to the last cob listed in 
Table 1. This was very small with a length of 2.6 cm., 
a diameter of 1.6 cm., and 12 kernel rows. The senior 
author, having shortly before studying the Chilean speci- 

TABLE 1 


Characteristics of the cobs from the earliest site (RAnL 100), 
dated at A.D. 105. 


diam/length 


code no. length(cem) diameter (cm) ratio kernel-row no. 

100-2 Sit 1.8 AQ 22 
100-3 3.6 Led 31 14 
7 3.1 1.3 42 18 
3.4 1.4 Al 16 

7 2.1 1.5 72 16 
100-7 6.3 1.8 .29 18 
6.2 1.6 .26 12 

7 6.1 1.7 .28 18 

rs 5.3 1.1 21 12 
100-4 5.6 2.1 37 18 
5.4 1.3 24 16 

? 57 1.6 .28 16 
5.1 1.5 .29 16 

100-5 7.0 2.8 40 30 
: 5.2 2.0 38 18 
4.2 1.6 38 16 

4.1 1.9 46 22 

“ 3.6 2.0 56 18 

7 3.6 1:5. 42 16 

“ 2.6 0.8 31 12 

average 4.6 1.6 35 17.2 


mens been engaged in studies of the prehistoric wild corn 
of the Tehuacin Valley in Mexico, saw in this specimen 
some resemblance to the Mexican corn; this raised in 
his mind the question of whether there could once have 
been a wild corn in Chile. Subsequent correspondence 
with the Junior author made it clear that this part of 
Chile could scarcely have provided a suitable habitat for 
wild corn, and we concluded that this specimen is proba- 
bly that of a stunted ear, borne on a depauperate plant. 

K.xcept for their smaller size, these earlier Chilean cobs 
(see Plate NV, fig. A) are quite similar in their charac- 
teristics to cobs of the modern race Capio Chico. The 
average data for their lengths, diameters, diameter/ 
length ratios, and kernel-row numbers are 4.6, 1.6, 0.35, 
and 17.2 respectively. For their modern counterparts, 
the corresponding averages, published by Timothy et a/., 
are 8.6, 2.38, 0.27, and 17.2 respectively. The similarity 
in kernel-row numbers is especially significant. 

In their dimensions, the Chilean cobs resemble even 
more closely the prehistoric ears excavated by Dr. 
Dwight Wallace from the Los Cerillos site in the Ica 
Valley on the south coast of Peru, some 1100 km. north 
of the Chilean site. This corn, estimated to be 2300- 
2500 years old, has been briefly described by Grobman 
et al, (1961) as a prehistoric race, Confite [queno; three 
ears of this race are illustrated in their figure 19. 

The specimens from the Los Cerillos site are unusually 
well preserved, the kernels still being attached to their 
cobs. To obtain estimates of the diameters of the cobs, 
we subtracted from the diameters of the ears 9/10 of the 
length of two average kernels, assuming that 1/10 of 
their length, on the average, was embedded in the cob; 
this figure is based on the data published by ‘Timothy 
et al. tor Capio Chico. 

Kistimates obtained in this manner from the Los Ceril- 


los maize produced the following averages: 5.6, 1.8, 
0.32, and 17.8. These are so similar to those set forth 
above for the prehistoric Chilean cobs that we may con- 
clude, with some degree of confidence, that the two 
collections represent essentially the same race, Confite 
Iqueno. By the same token, we can assume that if the 
Chilean cobs had retained their kernels, the intact ears 
would have been similar in their general appearance to 
those of the Los Cerillos maize, illustrated by Grobman 
et al. in their figure 19. 


Polulo 


This race is represented among the oldest cobs by a 
single specimen which is so different from the remaining 
ones that it must be considered as that of a different 
race. This specimen is 3.4 cm. long, but it is probably 
not intact with respect to length, since it lacks a pedun- 
cle. Also, since it comprises only the rachis, the central 
stem of the cob, the floral bracts having been lost, its 
diameter/length is not comparable with other cobs in 
this collection. The kernel-row number is 10; the cu- 
pules are distinct, slightly longer than wide and are 
hairy; stumps of the rachillae are prominent. 

We might not have been able to identify this single 
specimen had not we found counterparts of it in a collec- 
tion from site RAnL 887-1. Cobs of that lot, illustrated 
in Plate XV, fig. B, were derived from the shallow fill 
within the foundation of an isolated house dating to the 
late prehispanic period. 

Anticipating a description of this lot, to be set forth 
later, we can say that of the 115 cobs of site 8387-1, 
thirty-five are of a very distinctive type. These have 
slender rachises and peduncles of about the same diame- 
ter as the rachises. The floral bracts that remain attached 
to the rachises are relatively long; both lower and upper 


Loe | 


EXPLANATION OF THE PLATE 


Fic. A. Three of the larger cobs from lot 100, dated at ca. A.D. 
100. These are quite similar in the average dimensions to the pre- 
historic Peruvian race Confite Iqueno, and are related to the modern 
Chilean race Capio Chico Chileno. (Actual size. ) 


Fic. B. Three of the more slender cobs from lot 337-1, dated at 
ca. A.D. 1250. These are related to the modern Chilean race Polu- 
lo, which may in turn be related to the Peruvian popcorn Confite 


Morocho. (Actual size. ) 


Fic. C. Two cobs from lot 290, dating to the late prehispanic period. 
These are prehistoric counterparts of the modern Chilean race Chutu- 
cuno Chico, which may be related to the Peruvian race Confite Puntia- 
gudo. Note the stiff, curved lower glumes, a characteristic which may 
have been derived from one of corn’s relatives, teosinte or 7'ripsacum. 


(Actual size. ) 


[ 54 | 


AX “LIV Td 


glumes are herbaceous and not at all indurated. They 
are typical of certain types of pod-corn involving lower 
alleles at the pod-corn locus in combination with 7%, a 
major tunicate-inhibiting gene (see Mangelsdorf 1974). 

There is only one known modern Chilean race, Polulo, 
to which these slender cobs can be assigned. This is a 
finger-shaped popcorn with 10-16 kernel rows, grown at 
approximately 2700 meters. Its glumes are soft, the 
rachillae long and slender. 

Timothy et a/. state that the race Polulo seems not to 
be duplicated among collections made in other South 
American countries, but there seems to us a possibility 
that it is related to the Peruvian popcorn race Confite 
Morocho. Indeed one of the ears of Polulo illustrated 
in figure 5 of Timothy et al. resembles rather closely one 
of the ears of Confite Morocho illustrated in figure 48 of 
Grobman et a/. (1961). Both have about the same length, 
9.1 and 8.5 cm. respectively; both have 10 kernel rows 
and their rows are slightly irregular. One of the most 
distinctive characteristics of the archaeological specimens 
is their peduncles, which have about the same diameters 
as the rachises. The data of Grobman ect al. show that 
this is true of Confite Morocho. 

Galinat (1969) has isolated from the Peruvian race 
Confite Morocho types with cobs so slender that he calls 
them ‘‘string cobs.” These slender cobs are found also 
in the Peruvian race Rabo de Zorro, which Grobman 
et al, regard as a hybrid derivative of Confite Morocho. 
Moulds thought to represent ears of Rabo de Zorro occur 
on a number of ceramic vessels of the Moche culture 
(Early Intermediate Period); several of these are illus- 
trated in Grobman et al. (figs. 81 and 82), and one in 
Mangelsdorf (1974, fig. 17.18). These show that this 
slender-cob trait in South American races is an ancient 
one. The fact that it is not known in Mexican races may 


[ 57 


lend some support to the hypothesis (Mangelsdorf 1974) 
that there may have been an independent domestication 
of maize in South America. [t is in this connection that 
the slender-cobbed Chilean specimens are of particular 
interest. 

Galinat (1972) has recently published a brief descrip- 
tion of the archaeological maize turned up by MacNeish 
in asitein Ayacucho, Peru. He states that some of the 
cobs resemble those of the race Confite Morocho, but 
these are not among the earliest cobs from this. site. 
They are tentatively dated at 8000 B.P. 


Chutucuno Chico 


This race, described by Timothy ef af, appears to be 
represented by one specimen obtained from near the top 
of looters’ backdirt adjacent to the cemetery on site 100. 


~ 


This single cob is 7.6 em. long: its most prominent 
feature is the stiff indurated lower glumes. 

This specimen could be a precursor of cobs with stiff 
indurated lower glumes occurring in lots from. sites 
RAnL 186, 268B, and 290, described briefly below. 
These in turn appear to be related to the living Chilean 
race Chutucuno Chico which is grown at altitudes of 
2260 to 2500 meters. Timothy et al. describe this race 
as a small yellow popcorn with large cobs having 16-22 
or more kernel rows. Both red and white cobs occur in 
the modern as well as the late prehistoric collections. 

Chutucuno is in some respects similar to the fasciated 
form of the Peruvian primitive race Confite Puntiagudo 
described by Grobman ef a/. and illustrated in their fig. 
56. These authors attribute the indurated tissues of the 
rachis and lower glumes to hybridization with corn’s 
relative T'ripsacum. 


Maize Frou LATER Sires 
RAnL 2 
[ 58 | 


This site is a cemetery of more than 100 niche and pit 
graves at the locality of Chiu Chiu, and appears to have 
been utilized ca. A.D. 800. All the burials were found 
to have been looted, with broken artifacts and organic 
remains, including maize cobs, strewn on the ground 
surface. 

Fifty intact or almost intact cobs and six fragments 
were collected from the surface scatter. Of the intact 
cobs, twenty-nine are red and twenty-one white. Many 
are quite tripsacoid in having indurated tissues of the 
rachis and glumes. These cobs are quite similar to those 
of the lot from site RAnL 290 except that they are 
smaller. In shape they are tapered and rounded at the 
butts. The shanks are intermediate in thickness; the 
lower glumes are stiff, and the rachillae long. Measure- 
ments of five red and five white cobs show no significant 
differences in the two types except perhaps in kernel row 
numbers. The averages for lengths, diameters, and kernel 
row numbers are 7.6 cm., 1.7 ¢m., and 18.0 for the red 
cobs, and 7.5 cm., 1.6 em., and 15.6 for the white. 


RAnL 1 


This is a village site located at the present dispersed 
settlement of Lasana beside the Loa River, and may 
have been occupied from ca. A.D. 800 until Spanish 
arrival in the early 16th century. Maize cobs were col- 
lected from scattered surface refuse, and comprise eleven 
intact or almost intact specimens and six fragments. 

Three of the specimens resemble the slender cobs in 
the lot from site 837-1, and three resemble the thicker 
cobs from that same site. Five cobs are quite tripsacoid, 
with their tissues highly indurated either naturally or 
hardened through some kind of impregnation. One of 
the cobs has several single spikelets, a characteristic of 
corn’s relatives teosinte and T7ipsacum; both of the lat- 


[ 59 ] 


ter differ from corn in having solitary instead of paired 
pistillate spikelets. Three of the cobs have stumps of 
what may have been staminate spikes, 


RAnL 290 


This is a small burial site of niche and shaft graves 
near Chiu Chiu, dating to the late prehispanic period. 
All graves were found looted, but a collection of maize 
cobs was made from the surface of the burial chambers. 

Thirty-seven cobs were collected, of which eighteen 
are red or pink, seven variegated, and twelve white. The 
predominating shape is tapering with rounded butts: the 
shanks are intermediate in thickness. The lower glumes 
are stiff, indurated and glabrous. On several of the cobs 
some of the lower glumes have spots of brown pigmenta- 
tion characteristic of maize-teosinte or maize- T’ripsacum 
hybrids. The rachillae are prominent and in some speci- 
mens protrude beyond the lower glumes. 

There are no marked differences between the red cobs 
and the white. Averages for lengths, diameters, and 
kernel row numbers for eight red cobs are 9.0, 2.4, and 
18.3; for six white cobs these measurements are 8.9, 
2.6, and 18.3. ‘Two of the red cobs from this site are 
illustrated in Plate NV, fig. C. Most of the cobs appear 
to be related to the modern race Chutucuno Chico, which 
still occurs in Antofagasta Province. 


RAnL 186 


Dating to the late prehispanic period, this site consists 
of a single isolated shelter/dwelling situated on barren 
terrain 2 km. from the Loa River. A cache of between 
400 and 500 maize cobs was found in the earth floor of 
the structure. 

The cobs are quite variable in shape and other charac- 
teristics. Some are tapering at both ends; others are 


' 60 | 


more slender and more nearly cylindrical. With respect 
to diameter/length ratios, the cobs in a selected sample 
of twenty specimens vary from 0.13 to 0.86. In kernel- 
row numbers these same specimens vary from ten to 
twenty-six. Many of the cobs are twisted and contorted, 
and some are fused into conglomerate masses. ‘They must 
have been somewhat soft and pliable at one time, perhaps 
as the result of some chemical action in the soil. 

With respect to race, these cobs appear to represent 
a kind of ‘‘melting pot™’ in which the principal partici- 
pants were the races Capio Chico and Polulo; a minor 
component may have been Chutucuno Chico. 


RA nN iL 268B 


This is another small, looted cemetery of niche graves 
near Chiu Chiu, dating to the late prehispanic period. 
A cache of fifty-three cobs was found within one of the 
burial chambers. 

The specimens are similar to the cobs from site 290 
with respect to color; twenty-nine are red or pink, six 
variegated, and eighteen white, but are brighter due to 
better preservation. Another similarity is the stiff lower 
glumes, which in some specimens are curved like the 
teeth of a wood rasp. Most of the cobs appear to be re- 
lated to the modern Chilean race Chutucuno Chico. 


RAnL 337-1 


This isolated house site is located beside the Loa River 
among scrub vegetation, and is at least 12 km. from the 
nearest prehispanic settlement. The remains consist of a 
surtace scatter of artifacts and a 3.0 by 8.5 meter house 
foundation which contained a maximum of 28 cm. of 
unstratified refuse. A sample of wood from the excava- 
tion yielded a radiocarbon date of A.D. 1250 + 90 years 
(1-5899), thus placing the site within the late prehispanic 
period. 


[ 61 ] 


In one respect, the maize from the refuse is the most 
interesting of all of the lots recovered because it contains 
cobs of a slender type not previously described in the 
archaeological record of South America. 

The lot comprises 115 cobs, of which fifty are rather 
thick, tapering at both ends. The diameter/length ratios 
of these vary from 0.25 to 0.82 in five typical cobs that 
were measured; the kernel-row number varied from 
fourteen to eighteen. These cobs appear to be related to 
the modern Chilean race Capio Chico Chileno. 

The slender cobs (see Plate XV, fig. B) are thirty-five 
in number. In five cobs measured, diameter/length ratios 
varied from 0.09 to 0.14, and kernel row numbers from 
twelve to sixteen. Both upper and lower glumes are her- 
baceous, quite different from the stiff, indurated glumes 
of Chutucuno Chico, and they probably represent a form 
of pod-corn. As mentioned earlier in this report, they 
appear to be related to the Chilean race Polulo, which 
in turn may be a descendant of the primitive popcorn 
race Confite Morocho. 

Four kernels were found in this collection. All are 
popcorn: three are yellow in color and one is brown. 


Discussion 


The cobs of these collections, the earliest as well as the 
more recent, appear to be related to three still living 
Chilean races: Capio Chico Chileno, Polulo, and Chutu- 
cuno Chico. These in turn are related respectively to 
three prehistoric Peruvian races: Confite Iqueno, the 
popcorn race Confite Morocho, and the popcorn race 
Confite Puntiagudo. The last-named race resembles in 
a number of characteristics the Mexican popcorn Polo- 
mero ‘Toluqueno, but the remaining two races have no 
close counterparts in Mexican maize, either prehistoric 
or modern. This fact is consistent with the suggestion 


[ 62 | 


(see Mangelsdort 1974) that there may have been an in- 
dependent domestication of corn in South America. 


ACKNOWLEDGMENTS 


The field research during which the maize specimens 
were obtained was supported by a grant from. the 
National Science Foundation. The research was part of 
a Columbia University archaeological project in conjunc- 
tion with the Universidad del Norte in Antofagasta. 


| 63 | 


LITERATURE CITED 


Galinat, W.C. 1969. The evolution under domestication of the maize 
ear: string cob maize. Univ. Mass. Expt. Sta. Bull. 577. 


——, 1972. Common ancestry of the primitive races of maize indi- 
genous to the Ayacucho area in Peru. Maize Gen. Coop. News 
letter 46: 108-109. 


Grobman, A.; W. Salhuana, and R. Sevilla, in collaboration with 
P.C. Mangelsdorf. 1961. Races of maize in Peru. Nat. Acad. Sci.- 
Nat. Res. Council, Publ. 915. 


Mangelsdorf, P.C. 1974. Corn, its origin, evolution, and improve- 
ment. Harvard Univ. Press, Cambridge. 


Pollard, G.C. 1971. Cultural change and adaptation in the central 
Atacama Desert of northern Chile. Nawpa Pacha 9: 41-64. 


Timothy, D.H.: B. Pefa V., and R. Ramirez E., in collaboration 
with W.L. Brown and E. Anderson. 1961. Races of maize in Chile. 
Nat. Acad. Sci.-Nat. Res. Council, Publ. 847. 


[ G4 | 


ger 10 19/5 


NICAL MUSEUM LEAFLETS 
HARVARD UNIVERSITY 


CamspripGr, Massacnusetts, JUNE 16, 1975 Vor. 24, No. 4 


THE CONTRIBUTION OF LINNAEUS 
TO ORCHIDOLOGY'! 
BY 
Wintuiam T. STearNn? 


The presentation to the Botanical Museum, Harvard 
University of a collection of works by Carl Linnaeus 
(1707-1778), which formed part of the library of Oakes 
Ames (1874-1950), provides a fitting occasion on which 
to review concisely Linnaeus’s pioneer contribution to 
orchidology. Orchids were Ames’s life-study and, while 
himself contributing much new knowledge, he was too 
wise and well-informed to ignore the historic background 
of modern work; the scholarly essays gathered together 
in his Orchids in Retrospect (1948) manifest the range of 
his orchidological learning and contain many references 
to earlier literature. 

Orchidology is the study of orchids as a special group 
within the Plant Kingdom. Its existence necessarily de- 
pends upon recognition of them as a definable group 
worthy of particular study, and that implies a high degree 
of botanical sophistication. A group embracing plants so 

'This paper embodies a lecture given at the Botanical Museum of 
Harvard University on September 18, 1969 when the heirs of Mrs. 
Blanche Ames presented Professor Oakes Ames’s collection of Lin- 
naeana to the Botanical Museum. It includes part of an earlier lec- 
ture, “I'wo thousand years of orchidology’, printed in Proc. Third 
World Orchid Conf. 1960, 26-42 (1960). 

* Department of Botany, British Museum (Natural History), Lon- 


don, England. 
5 


[ 65 ] 


ECONOMIC BOTANY LIBRARY 
OF OAKES AMES 
HARVARD BOTANICAL MUSEUM 


distinct in habit as Hria microphylla, with leaves only 1 
cm. long, creeping over the bark of tropical trees, and the 
tall erect terrestrial savanna herbs of the genus Hudophia, 
as well as the saprophytes exemplified by phyllorchis, 
Corallorhiza, Didymoplexis, E:pipogium, Gastrodia and 
Neottia, which spend most of their lives hidden in sub- 
terranean darkness, can be held together in a single bo- 
tanical family, the Orchidaceae, only by emphasis on a 
common basic floral pattern by no means obvious to the 
uninitiated. The existence of such a ground plan of floral 
organization tends to be masked by fantastic variations 
upon it in size, shape and colour associated with pollina- 
tion mechanisms so complex as to be unbelievable were 
they not so well attested. Consequently, the concept of 
the Orchidaceae as a group of petaloid monocotyledons 
technically distinguished by such associated characters 
as a perigon of six segments zygomorphically developed, 
stamens reduced to one, two or three, ovary inferior and 
usually twisted (the whole flower then upside down) a 
stigma modified into a peculiar organ, the rostellum, 
seeds very numerous, minute and lacking endosperm— 
this concept has been long in the making. 

The orchids lack a single marked feature comparable 
to the umbel of Umbelliferae (Apiaceae), which led to the 
‘arly recognition of that aromatic economically impor- 
tant family. Nevertheless, they now present themselves 
to us as a natural group without conscious reference to 
their individual technical features. This recognition is 
the result of familiarity. The firm waxy texture of their 
flowers, in shape generally unlike those of other plants, 
their frequent use in floristry and their former association 
with wealth have established orchids as a generally known 
and popular group. They continually provide botanists 
with new problems and gardeners with new joys un- 
paralleled in any other botanical family. 


| 66 | 


When discussing the evolution of the orchid flower in 
Amer. Orchid Soc. Bull. 15: 18-19 (1946), Ames re- 
marked, with light-hearted scholarship, that ‘certain 
floral structures and the story of their probable evolution 
intensify the belief that orchids possess qualities which 
set them apart from other plants. Therefore (in the bosom 
of the family, with no botanists present) it is delightful to 
regard orchids as beings which have shaped their own des- 
tiny and to speak of their floral structures, of the rostel- 
lum, for example, as organs purposefully developed to 
serve special and fantastic functions’. Inthe 18th century, 
Linnaeus recognized most of their distinctive features asa 
group, although he knew only a few species. He thus pro- 
vided a base upon which, in the 19th century, other bota- 
nists, John Lindley their chief, built modern orchidology. 

The words orchis (6pxes) ‘testicle’ and logos (Aoyos) 
‘discourse’ are Greek, but the distinctive technical details 
of the family Orchidaceae if made known to a learned and 
intelligent Ancient Greek herbalist, even Dioscorides 
himself, in the Ist century A.D. would have had little 
or no meaning or interest. ‘To people whose interest in 
plants was primarily utilitarian, the division of the Plant 
Kingdom into major groups defined by inconspicuous and 
apparently trivial features such as these floral subtleties 
of organization—groups which Magnol called familae 
and Linnaeus ordines naturales—would have seemed 
pointless. The Ancient Greeks and Romans, like their 
successors in Kurope down to the 17th century, knew 
nothing about the functions of the different parts of the 
flower; they lacked lenses suitable for supplementing and 
training the naked eye in observation; consequently, 
they gave little attention to the floral structure on which 
the concept of the Orchidaceae rests. For them the word 
orchiologos would have suggested either a medical treatise 
or an obscene comedy. 


[ 67 | 


To survive the long, dry, fiercely hot Mediterranean 
summer, plants must either have features enabling them 
to withstand dessication, such as small, hard or woolly 
leaves, or else disappear underground, having stored 
during the relatively mild moist winter and spring 
enough food in bulbs or tubers to give them a quick 
start into growth when the rains come. The storage 
organs of a number of Mediterranean orchids such as 
Orchis and Ophrys are paired rounded tubers which sug- 
gested testicles to the far from prudish Greeks, and they 
named these plants orchis (dpyes) and cynosorchis 
(kuvvocopxts) ‘dog’s testicle’ accordingly, a matter dis- 
cussed by Ames in his little essay on the ‘Origin of the 
term Orchis’ in Amer. Orchid Soc. Bull. 11: 146-147 
(1942). Dioscorides described the root as bulbous, with 
paired swellings, one full and solid, the other wrinkled 
and soft, and mentioned asa folk-tale the belief that men 
should eat the full one to beget sons but women the soft 
one to conceive daughters, presumably avoiding syn- 
chronisation! The extension of the word orchis and cog- 
nate words to tropical plants with long slender dangling 
roots would have seemed an absurdity for them, a con- 
tradiction in terms. The extension of the concept of 
‘orchid’ to cover such plants owes much to Carl Linn- 
aeus (after 1762 Carl von Linné) but did not originate 
with him. It developed out of the studies of his predeces- 
sors who had a feeling for affinities between plants as ex- 
pressed in overall resemblance. 

Thus Gaspard (or Caspar) Bauhin (1560-1624), who 
attempted in his Prnav (1628) to regulate the names of 
all the plants then known, devoted section 6 of his book 
Il mainly to orchids, with headings now unfamiliar such 
as Cynosorchis seu Testiculus canis, Orchis Serapias, ete., 
although somewhat spoiling the picture by associating 
Dens canis (ce. Hrythronium) and Orobanche with them. 


| 68 | 


J 


PuatTE XVI 


TRIORCHIS SERA Knabentrautmennle das tleines, 


PIAS MAS, 
Orchis morio L., illustrating the paired tubers which suggested the 
generic name. Reproduced from Leonhard Fuchs, De historia stirpium, 
page 559, 1542, Basel. 


[ 69 ] 


Towards the end of the 17th century came the huge 
works of John Ray (1628-1705) and Robert Morison 
(1620-1683). Ray’s account occupies pages 1212-12338 
of volume 2 of his Historia Plantarum (1688), the spe- 
cies being listed under the names Orchis, Tragorchis, 
Cynosorchis, Chamaeorchis, Monorchis, Triorchis, Palm- 
ata, Pseudo-Orchis, Nidus-avis, Pscudo-limodoron, Helle- 
borine and Bifolium. He possessed a good knowledge of 
these plants derived from his extensive travels in Britain 
and on the Continent of Kurope, and his descriptions 
give significant details as regards the species but no for- 
mal generic characters. His comments on the great difh- 
culty of studying orchids on account of the numerous 
allied species and the inadequate descriptions by bota- 
nists, of which one description can apply to several spe- 
cies and several different descriptions to one species, has 
unfortunately not ceased to be true: ‘Caeterum cum 
Orchidum historia magna confusione et obscuritate lab- 
oret, partim ob multitudinem specierum, et nonnullarum 
etiam similitudinem, partim ob earum apud_ botanicos 
descriptiones, adeo breves interdum et generales ut et 
una pluribus, et plures uni alicul speciel accommodari 
possint: unde perspicacissimum quemque et in his studiis 
versatissimum torqueant et perplexum reddant necesse 
est’ (ist. 2: 1212: 1685). This was a state of affairs 
which Linnaeus set out to remedy. 

In the third volume (1699) of Morison’s Plantarum 
FAistoria universalis Oxvoniensis, which was completed by 
Jacob Bobart after Morison’s death, 108 kinds of Orchid- 
aceae are listed together, though associated with other 
plants having minute seeds such as Orobanche and Pyr- 
ola. These are put in six genera: Helleborine, Bifolium 
sive Ophris, Triorchis et Monorchis, Orchis palmata, 
Pseud-orchis. 

Joseph Pitton de Tournefort (1656-1708) in his In- 


[ 71 ] 


stitutiones Rei Herbariae 1: 481-488 (1701) dealt with 
them as a special group under the heading ‘De Herbis, 
flore polypetalo, anomalo’, this including Orchis, Helle- 
borine, Calceolus (i.e. Cypripedium), Ophris (i.e. Listera), 
Limodorum and Nidus avis (i.e. Neottia). Unlike his 
predecessors, he provided concise generic descriptions. 
Within Orehis he included plants with flowers resembling 
‘nunc Hominem nudum, modo Papilionem, Fucum, 
Columbam, Simiam, Lacertam, Psittacum, Muscam, 
caeterave repraesentante’ now put in Orchis, Dactylo- 
rhiza, Ophrys, Himantoglossum, Aceras, ete. This ex- 
traordinary mimicry of animal forms by the flowers of 
orchids had already excited the admiration of Jacob 
Breyne (1637-1697) in his Hwvoticarum aliorumque minus 
cognitarum Plantarum, 94 (1678) as Ames noted in his 
essay, ‘Orchids in retrospect’ (in Amer. Orchid Soc. Bull. 
11: 102-106; 1942). The orchids known to these early 
authors were north-temperate species. Of the astonish- 
ing richness of the tropics in orchids they knew nothing. 

In 1708, H.A. van Rheede tot Draakestein (1685 
1691) posthumously recorded six orchids from Malabar, 
southern India, in his Hortus Indicus Malabaricus, these 
being now known as Rhynchostylis retusa, Vanda spath- 
ulata, Acampe Wightiana, Sarcanthus peninsularis, Den- 
drobium ovatum and Cymbidium alotfolium. 

On his visit to Jamaica in 1687 to 1689, Sir Hans 
Sloane (1660-1753) found a number of epiphytic orchids. 
Those recorded in his Voyage to the Islands Madera xxx 
and Jamaica (1707-25) as species of Viscum, Cardamon, 
ete. are now known as Brassavola cordata, Oncidium 
guttatum, Broughtonia sanguinea, Vanilla claviculata, 
Stenorrhynchus speciosus and Hrythrodes plantaginea. 

About the same time, another doctor, Engelbert 
Kaempfer (1651-1716), was becoming acquainted with 
a few of the many orchids of Java and Japan. He illus- 


> 


[ 72] 


trated three of these in his Amoenitatum Exvoticarum 
Fasciculi V (1712), the species now known as Dendrobium 
moniliforme, Vanilla domestica and Arachnis flos-aeris. 
Such was the state of knowledge of orchids at and 
after the birth of Carl Linnaeus in 1707. His career has 
been sketched in so many publications, notably B.D. 
Jackson, Linnaeus (afterwards Carl von Linné), the 
Story of his Life (1923), Kk. Hagberg, Carl Linnaeus 
(1952), N. Gourlie, The Prince of Botanists (19538), and 
W. Blunt and W.'T. Stearn, The Compleat Naturalist 
(1971), that a bare outline will suffice here. He was born 
in 1707 at Rashult, Skane, southern Sweden, entered 
the University of Lund in 1727, changed next year to 
the University of Uppsala, made an important journey 
to Lapland in 1782, left Sweden for Holland in 1785 and 
returned in 1788, having acquired a doctor’s degree at 
Harderwijk and published his Systema Naturae (1735), 
Fundamenta botanica (1736), Genera Plantarum (1787), 
Flora Lapponica (1737), Critica botanica (1737), Hortus 
Chffortianus (1788). He was appointed a professor at the 
University of Uppsala in 1741 and that year visited the 
Baltic islands of Oland and Gotland. His Species Plan- 
tarum, the starting point of modern botanical nomen- 
clature, appeared in 1758, and his Systema Naturae, 10th 
ed., vol. i, the starting point of modern zoological no- 
menclature, in 1758. He died at the age of 70 in 1778. 
On his journey to Lapland in 17382, Linnaeus became 
acquainted with seven species now known as Dactylo- 
rhiza maculata, Coeloglossum viride, Goodyera repens, 
Corallorhiza trifida, Listera cordata, Herminium mon- 
orchis and Cypripedium calceolus, admittedly but not sur- 
prisingly few, considering their remoteness from the 
tropics, and also atypical of this essentially tropical 
family. He knew Calypso bulbosa only from a coloured 
drawing made in Lapland by Rudbeck the Younger. 


[ 73 


These species he enumerated in his Flora Lapponica 
(1737). 

In 1738, Linnaeus published his Classes Plantarum 
which summarizes the earlier classifications of the Vege- 
table Kingdom by his predecessors, Cesalpino, Morison, 
Ray, Knaut, Hermann, Boerhaave, Rivinus, Rupp, Lud- 
wig, Tournefort, Pontedera and Magnol, as well as his 
own artificial “sexual system’, wherein the genera now 
included in Orchidaceae form his class Gynandria order 
Diandria. He appended to this a ‘Fragmenta methodi 
naturalis’ which is an attempt at a more natural system 
of classification. Here again, as in his ‘systema sexuale’, 
the orchid genera, Orehis, Satyrium, Serapias, Hermi- 
nium, Neottia, Ophrys, Cypripedium and Epidendrum, 
form a group distinct from other plants, his Ordo IV. 
Much later, in his Genera Plantarum, 6th ed. (1764), 
he published a modified version under the heading 
‘Ordines naturales’; this is reprinted, with a discussion 
by Stearn, in the Ray Society facsimile of Iinnaeus, 
Species Plantarum, vol. 2, Appendix: 98-102 (1959). 
Here appears an Ordo VII Orchidaceae comprising the 
above genera. After his death, two former students, 
P. 1). Giseke and J.C. Fabricius, published a detailed 
account of lectures on the natural system delivered by 
Linnaeus at Uppsala in 1764 and 1771 entitled Caroli a 
Linne Praelectiones in) Ordines naturales Plantarum 
(1792). Here again the Orchideae from a distinct “Ordo™ 
(in modern terminology, family) of which Linnaeus 
stated the characters, the ecology and the cultivation, 
then so little understood that most tropical orchids died 
soon after their introduction. “Omnium difficillime in 
hortis plantantur, et introductae vix per aliquot annos 
perennant’, he said, ‘I seminibus satis in hortis non pro- 
veniunt: certe si possent seri, nullus hortulus amoeni- 
oribus floribus superbiret’. Their successful cultivation 


[ 74 J 


and raising from seed was not achieved until the 19th 
century; John Smith (1798-1888) provided in_ his 
Records of the Royal Botanic Gardens, Kew, 227-241 
(1880) a first-hand account of progress in orchid-growing 
during his lifetime. 

On his way back to Sweden from Holland in 1738, 
Linnaeus visited Paris and was conducted to Fontain- 
bleau, where the wild orchids in particular delighted him. 
Before taking up his professorship at Uppsala in October 
1741, Linnaeus with six young companions made an ex- 
pedition to Oland and Gotland at the request of the 
Swedish Estates of the Realm. The purpose of this Jour- 
ney, which extended from May to August 1741, was 
primarily economic, to search for dye-yielding and medi- 
cinal plants and to make observations on soils, minerals, 
farming, manufacturing methods and general natural 
history. These islands, then scientifically almost unex- 
plored, differ from mainland Sweden in being of lime- 
stone and having a relatively mild climate, so that they 
shelter many organisms rare or unknown on the main- 
land, among them various orchids. The two islands natu- 
rally excited Linnaeus’s eager curiosity, and he described 
them in detail in his entertaining account of the journey 
entitled Oliindsha och Gothlindska Resa (1745), of which 
an annotated English translation by Marie Asberg and 
W.'T. Stearn is now available in Biol. J. Linnean Soc. 
5: 1-220 (1973). Their orchid flora includes 22 species. 
Linnaeus visited the islands when most of the orchids 
were in full flower. As soon as he and his companions 
touched the shore of Oland on June 12 (June 1, Old 
Style, of Linnaeus’s journal) they realized that this was 
a land altogether different from the other Swedish prov- 
inces and they decided to make meticulous notes about 
everything. Their second day provided them with the 
joy of finding Ophrys insectifera, Orchis militaris, Orchis 


[ 75 | 


ustulata, Orchis morio, Gymnadenia conopsea, Dactylo- 
rhiza sambucina, Dactylorhiza maculata and Dactylorhiza 
incarnata and caused Linnaeus to remark that, such 
plants having previously been unrecorded for Sweden, 
he had never expected when seeing them at Fontainbleau 
that he would ever see them again. These grew between 
Forjestaden and Bornholm. Listera cordata was found 
on June 15 (June 4, Old Style). Later (June 25, Old 
Style), he came across Orehis mascula in Lummeland 
parish north of Visby; he also found Herminium mon- 
orchis (July 5, Old Style), Mpipactis helleborine (July 25, 
Old Style) and Mpipactis atrorubens (July 8, Old Style). 

These Baltic orchids inspired Linnaeus to give special 
attention to all of the species then known. Many had 
been described by earlier authors under a diversity of 
names. Linnaeus reduced them to 41 species in a sur- 
vey entitled ‘Species orchidum et affinitum plantarum’ in 
Acta Soc. Regiae Sei. Upsal. 1744: 1-87 (1744) cited 
by Linnaeus as ‘Act. ups. 1740°. For each one he gave 
very detailed synonymy and a concise diagnosis. There 
are no generic descriptions here, as he had already pro- 
vided them in his Genera Plantarum, 2nd edition (1742). 
His genera were few: Orchis (including Dactylorhiza), 
Satyrium, Serapias, Herminium, Neottia, Ophrys, Cy- 
pripedium and I/pidendrum. 

In 1745 Linnaeus published the account of his Oland 
and Gotland journey, Oltindska och Gothlindshka Resa, 
already mentioned. An important feature of this inter- 
esting book is its index because it foreshadows Linnaeus’s 
later introduction of consistent binomial nomenclature 
for all species of plants and animals. In the text he used 
diagnostic polynomials such as Orchis bulbis indivisis, 
nectaru labio quinquefido punctis scabro, cornu obtuso, 
petalis conniventibus and Orehis bulbis palmatis, nectari 
cornu setaceo germinibus longiore, labio crenato, names 


[ 76 | 


which indicate the distinguishing features of the species 
but are inconvenient to write and almost impossible to 
remember. In the index these are listed as Orchis 725 
militaris and Orchis 727 longicalcar, the number referring 
to the numbered entry with diagnostic details in Linn- 
aeus’s Hlora Suecica (1745). The generic name and the 
added single epithet were easy to remember; the num- 
ber took the place of the long descriptive phrase by serv- 
ing as a concise reference to one in a standard work. 
These names thus provide an intermediate stage between 
the polynomial nomenclature of Linnaeus’s Flora Sue- 
cica (1745) and the binomial nomenclature of his Species 
Plantarum (17538), where, for example, the two species 
mentioned above are simply designated Orchis militaris 
and Orchis conopsea; the last species, however, has long 
been removed from Orc/is and is now known as Gym- 
nadenia conopsea. Apart, therefore, from its interest as 
an 18th-century travel book replete with first hand ob- 
servations on local customs and superstitions—plants, 
animals, minerals, quarrying, household remedies, ver- 
nacular names, runic inscriptions and much else— Linn- 
acus’s Oland and Gotland Journey 1741 is an important 
work in the history of biology. 


Orchids for Linnaeus were only a small part of the 
Vegetable Kingdom which he had set himself the daunt- 
ing task of classifying and naming. He accomplished this 
in his Species Plantarum (1758) and the associated fitth 
edition (1754) of his Genera Plantarum. Contrary to 
common belief, the introduction of consistent binomial 
nomenclature for species, i.e. the designation of each one 
by a two-word name, e.g. Orchis ustulata, consisting of 
the generic name, e.g. Orchis, followed a single specific 
epithet, e.g. wstu/ata, was not the primary intent of the 
Species Plantarum, although this nomenclatural innova- 
tion has given it outstanding importance. The addition 


[77 | 


of specific epithets came almost as an after-thought: as 
late as 1748, Linnaeus had not added them to his manu- 
script draft now in the library of the Linnean Society of 
London. The main purpose of the Species Plantarum was 
to provide diagnostic phrase-names (polynomials), such as 
Cypripedium radicibus fibrosis, foltis ovato-lanceolatis cau- 
linis and Cypripedium bulbo subrotundo, folio subrotundo 
radical, which, taken together, functioned like the con- 
trasts in a modern key and enabled the species to be dis- 
tinguished from their congeners, with relevant synonyms 
cited as a guide to the existing literature about each. 
Hence, the material, either specimens or illustrations, sup- 
plying the information epitomised in the phrase-name, is 
of first importance for the typification of the associated 
specific epithet. Hach phrase-name set out to give as con- 
cisely as possible a statement of the main distinguishing 
features. Thus, in his synopsis of the genus Orc/is in the 
Species Plantarum (1: 989-944), the phrase-names of spe- 
cies 1-10 begin with ‘bulbis indivisis’ (‘with undivided 
root-tubers’) and so distinguish them from species 11-14 
with ‘bulbis palmatis’ (‘with palmately lobed root-tubers’ ) 
as well as from species 15-17 with ‘bulbis fasciculatis’ 
(‘with bunched fibrous roots’). A number of species are 
further distinguished by the length of the spur in relation 
to the ovary, e.g. species 12. (maculata) with ‘nectarii 
cornu germinibus breviore’ (‘with spur shorter than the 
ovary ) being thereby distinguished from species 18 
(conopsea) with ‘nectaril cornu setaceo germinibus lon- 
giore’ (‘with spur setaceous longer than the ovary’) and 
species 14 (flava) with ‘nectarii cornu filiformi longitu- 
dine germinis’ (‘with spur filiform as long as the ovary’). 
A name such as that for species 5 (pyramidalis), Orchis 
bulbis indivisis, nectaru labio trifido antice bidentato, cornu 
longo, petals acuminatis served to distinguish this spe- 
cies from all other orchids then known. 


[ 78 | 


Descriptions of the genera were provided in the fifth 
edition of the Genera Plantarum. Here, Linnaeus recog- 
nized as distinet Orchis (which included Tournefort’s 
Limodorum), Ophrys, Serapias (which included 'Tourne- 
fort’s Helleborine), Limodorum, Arethusa, Cypripedium 
(which included Tournefort’s Calceolus) and I/pidendrum 
(which included Plumier’s V’ani//a). The content of these 
genera can be best illustrated by listing the species in- 
cluded in each genus by Linnaeus in the Species Plant- 
arum and then giving the currently accepted name for 
each species and also the name which Oakes Ames 
adopted or would probably have adopted when dealing 
with the same species. This list, compiled with the help 
of Mr. Peter Fk. Hunt, is as follows: 


Linnaeus, Species Plantarum 2: 989-954 (1752) 


oe WwW bd ee 


-p OOOND 


tp 


12 
13 


14 
15 
16 
17 


Linnaean name 


Currently accepted name 
(if different from Linnaean name) 


Probable name used by Ames 
(if different from 


currently accepted name) 


Orchis susannae Pecteilis susannae (L.) Raf. Habenaria susannae (L.) 
R.Br. 
O. ciliaris Blephariglottis ciliaris (L.) Rydb. H. ciliaris (L.) R.Br. 
O. bifolia Platanthera bifolia (L.) L.C.Rich. H. bifolia (L.) R.Br. 
O. cucullata Neottianthe cucullata (L.) Schltr. H. cucullata (L.) Hoefft. 
O. pyramidalis Anacamptis pyramidalis (L.) 
L.C.Rich. 7 
O. coriophora — — 
O. cubitalis Peristylus cubitalis (L.) Kraenzlin H. cubitalis (L.) R.Br. 
O. morio _ — 
O. ustulata _— — 
O. militaris — a 
O. latifolia Dactylorhiza latifolia (L.) Soé O. latifolia L. 
(D. majalis (Rchb.) P.F.Hunt & 
Summerh. ) 
O. maculata D. maculata (L.) So6 O. maculata L. 
O. conopsea Gymnadenia conopsea (L.) R.Br. Habenaria conopsea (L.) 
Benth. (non Rchb.f.) 
O. flava Tulotis flava (L.) Sengh. H. flava (L.) R.Br. 
O. fuscescens Tulotis fuscescens (L.) Raf. H. fuscescens (L.) Torr. 
O. strateumatica Zeuxine strateumatica (L.) Schltr. — 
O. abortiva Limodorum abortivum (L.) Swartz — 


[79 ] 


Linnaeus, Species Plantarum (cont. ) 


Currently accepted name 
(if different from Linnaean name) 


Linnaean name Probable name used by Ames 
(if different from 
currently accepted name) 
Blephariglottis psycodes (L.) H. psycodes (L.) Sprengel 

Rydb. 


Galearis spectabilis (L.) Raf. 


18 O. psycodes 


19 O. spectabilis Orchis spectabilis L. 


Koch _ 
2S. viride Coeloglossum viride (L.) Hartman H. viridis (L.) R.Br. 
3. S. nigrum Nigritella nigra (L.) Rchb. — 
4 §S. albidum Pseudorchis albida (L.) Love & H. albida (L.) R.Br. 


Satyrium hircinum 


Himantoglossum hircinum (L.) 


Love 


5 S. epipogium Epipogium aphyllum Swartz — 
6 S. repens Goodyera repens (L.) R.Br. = 
1 Ophrys nidus-avis Neottia nidus-avis (L.) L.C.Rich. — 
2  O. corallorhiza Corallorhiza trifida Chatelain — 
3 O.spiralis Spiranthes spiralis (L.) Chevall. — 
4 ©. cernua S. cernua (L.) L.C.Rich. — 
5 O. ovata Listera ovata (L.) R.Br. —_ 
6 QO. cordata L. cordata (L.) R.Br. —_— 
7 O. lilifolia Liparis lilifolia (L.) L.C.Rich. 


8 O. loeselil L. loeselii (L.) L.C.Rich. — 
9 QO. paludosa Malaxis paludosa (L.) Swartz — 
10 O. monophyllos M. monophyllos (L.) Swartz — 
11. O. monorchis Herminium monorchis (L.) R.Br. a 
(‘Monochris’ ) 
12 O. alpina Chamorchis alpina (L.) L.C.Rich. — 
13. O. camtschatea Neottia camtschatea (L.) Rchb.f. — 
14 O. anthropophora Aceras anthropophorum (L.) R.Br. — 
15 O. insectifera —  (O. muscifera Hudson) — 
1 Serapias hellecorine Epipactis helleborine (L.) Crantz — 
latifolia “ (E. latifolia (L.) All. —_ 
longifolia Cephalanthera longifolia (L.) 
Fritsch _ 
palustris Epipactis palustris (L.) Crantz — 
2 S. lingua = — 
1 Limodorum tuberosum Bletia tuberosa (L.) Ames — 
1 Arethusa bulbosa — — 
2 A. ophioglossoides Pogonia ophioglossoides (L.) 
Ker-Gawl. —_— 
3. A. divaricata Cleistes divaricata (L.) Ames —_ 
1 Cypripedium calceolus — — 
2 C. bulbosum Calypso bulbosa (L.) Oakes — 
1 Epidendrum vanilla Vanilla planifolia Andrews Vanilla fragrans Ames 
2 E. domesticum Vanilla domestica (L.) Druce — 


ex Lindl. 


' 80 | 


Linnaeus, Species Plantarum (cont. ) 


Linnaean name Currently accepted name Probable name used by Ames 
(if different from Linnaean name) (if different from 
currently accepted name) 


3. E. flos-aéris Arachnis flos-aéris (L.) Rchb.f. — 
4 E. tenuifolium Cleisostoma tenuifolium (L.) Garay — 
5 E. spathulatum Vanda spathulata (L.) Sprengel —_ 
6 E. ovatum Dendrobium ovatum (L.) Kraenzlin — 
7 E. nodosum Brassavola nodosa (L.) Lindl. — 
8 E. carinatum Dendrobium carinatum (L.) Willd. — 
9 E. aloifolium Cymbidium aloifolium (L.) Swartz — 
10 ~E. guttatum Oncidium guttatum (L.) Rchb.f. 
11 E. retusum Rhynchostylis retusa (L.) Blume — 
12. E. amabile Phalaenopsis amabilis (L.) Blume — 
13 E. ensifolium Cymbidium ensifolium (L.) Swartz — 
14. E. moniliforme Dendrobium moniliforme (L.) Swartz — 


From the above concordance, it will be evident that 
Linnaeus’s 19 species of Orchis can be referred to 15 
genera or at least 5 genera; his 6 species of Satyrium to 
6 genera or at least 5 genera; his 15 species of Ophrys 
to 10 genera; his two species of Serapias to 8 genera: 
his 8 species of Arethusa to 8 genera; his two species of 
Cypripedium to 2 genera; his 14 species of H/pidendrum 
to 10 genera. Thus, in a group of species where Linn- 
aeus saw only 7 genera, modern orchidologists see 50. 
This change of viewpoint has come largely from the in- 
troduction into gardens of allied species during the 19th 
century, a consequence both of Kuropean botanical ex- 
ploration outside Europe and of improved methods of 
cultivation, the two together providing living specimens 
for study and greatly stimulating this study. Of the 
species accepted by Linnaeus in 1758, Europe provided 
30, eastern North America 11, Indonesia and the Philip- 
pines 4, Siberia 4, India 6, tropical America 8, Ceylon 
2, China 1; only 4 were previously undescribed species. 
He had a first-hand knowledge of many European spe- 
cies in a living state, but his acquaintance with tropical 
orchids was meagre. He knew his Orchis cubitalis (Peri- 


[ 81 ] 


stylus cubitalis) and O.stateumatica (Zeuvine stateumatica) 
from Ceylonese specimens in Hermann’s herbarium, now 
in the British Museum (Natural History), his Z’piden- 
drum amabile (Phalaenopsis amabilis) and I. ensifolium 
(Cymbidium ensifolium) trom Chinese specimens collected 
by Osbeck and his 14. Vanilla (Vanilla planifolia) trom 
cultivated material and 24. nodosum (Brassavola nodosa) 
from cultivated material and illustrations, but the other 
9 he knew only from illustrations published by Rheede 
and Kaempfer. Wisely he kept these little-known species 
in a single genus, Mpidendrum; he could record their 
existence, give them names but add nothing to an under- 
standing of them. This had to wait for Olof Swartz 
(1760-1858), who studied many species while in the 
West Indies, Robert Brown (1778-1858), who studied 
many in Australia, Carl Ludwig Blume (1796-1862), 
who studied many others in Java, and John Lindley 
(1799-1865), who never visited the tropics but studied 
their products more conveniently in the glasshouses of 
Kngland and published some 120 currently accepted 
genera. 

Assessment of Linnaeus’s main publications relating 
to orchids depends upon one’s viewpoint and this is con- 
ditioned by both the knowledge and the scientific fash- 
ions of the time. If, forexample, we imagine Carl Linn- 
aeus as a Harvard student submitting his account of 
Orchidaceae to Professor Ames as a thesis, which ad- 
mittedly requires some imagination, we can be sure poor 
student Linnaeus would have been taken to task for the 
superficiality of his work, for ignoring so many minute 
floral details in his definition of genera, for failing to 
separate groups which, in the Professor’s eyes, were ob- 
viously distinct; Linnaeus would have been told to ex- 
amine their floral structure with greater care. On the 
other hand, if we imagine Oakes Ames as an Uppsala 


[ 82 | 


student making his work on orchidaceous genera known 
to Professor Linnaeus, we can also imagine the latter 
severely questioning the propriety of his taxonomic pro- 
cedure. Linnaeus could well have criticized him for 
ignoring the overall resemblance of his plants which 
justified keeping them together, for giving excessive 
importance to insignificant details and for revealing by 
his absurd breaking up of natural genera (although Ames 
in fact was more conservative in his approach than some 
of his contemporaries) a lack of understanding of the 
essentials in classification as then understood. 

Both professors would have been right within the con- 
text of their times. When the known species of a family 
are few, it is probably more convenient to keep them 
together within a few genera. When the same species 
have had added to them a host of others, it may be more 
convenient to divide them all among a much greater 
number of genera. Provided the facts about them are 
understood, it is often a matter of convenience or pre- 
vailing fashion or consistency of treatment in comparable 
situations, or even plain egotism, as to the taxonomic 
rank these groups should be assigned. 

The value of Linnaeus’s contributions to orchidology 
lies primarily not in his treating orchids as a special group 
but, on the contrary, in giving them the same nomen- 
clatural treatment as other plants in accordance with his 
methodology. The essentials of this were that: 


1) the genera should be named euphoniously and clear- 
ly, avoiding names which could be confused ; 

2) the generashould be defined according to a consistent 
formal plan, using a fixed terminology ; 

3) the species should be defined by key characters en- 
abling them to be distinguished and recognized ; 

4) the species should be given convenient binomial 
names. 

[ 83 | 


Such procedure has been fundamental to the develop- 
ment and expansion of systematic botany in general and 
to orchidology in particular, to the work of Oakes Ames 
no less than to his predecessors, from Linnaeus onwards; 
but it now seems so commonplace and self-evident that 
only by comparison with the work of Linnaeus’s prede- 
cessors does its originality become manifest. Linnaeus 
distinguished in the Species Plantarum (1758) only 7 
genera and 62 species out of the 700 or so genera and 
20,000 or so species of Orchidaceae now known and, by 
the end of his life, had become acquainted with only 107 
or so genera. Nevertheless, his treatment of these few 
exemplified the methods which, with due modification, 
have made possible the recording of the many. Thus, in 
the works of Linnaeus lay the potentialities of the studies 
in orchidology so honourably associated with Ames and 
the Botanical Museum of Harvard University. 


[ 84 ] 


ECONOMIC BOTANY LIBRARY 
OF OAKES AMES 
HARVARD BOTANICAL MUSEUM 


BOTANICAL MUSEUM LEAFLETS 


HARVARD UNIVERSITY 


Campripngr, Massacuuserts, Jung 27, 1975 Voi. 24, No. 


K'THNOBOTANY OF THE WESTERN 
TARAHUMARA OF CHIHUAHUA, 
MEXICO 


I. Nores ON THE GENUS Agave 
BY 
Roserr A. Byrg, Jr.', Don Burcess’, 
AND ALBINO Mares Trias’ 


INTRODUCTION 


The genus 4 gave, native to Mexico and southwestern 
United States, has played an important role in the ma- 
terial culture of the peoples who lived in its range. In 
the Greater Southwest, various species played a part in 
almost every aspect of aboriginal life (Castetter, Bell and 
Grove, 1988; Gentry, 1972: 9-11). Kven in industri- 
alized Mexico, the maguey plants are important sources 
of commercial fibre (sisal and henequén) and fermented 
beverages (pulque). 

In southwestern Chihuahua, several species of fave 
are still important to the Tarahumara Indians, although 
the intensity of some uses has decreased. From the first 
European observations to the present, numerous writers 

' Botanical Museum of Harvard University. 

* Missionary-linguist, Summer Institute of Linguistics, Rocoroibo, 
Municipio de Guazapares, Chihuahua. 

* A Western Tarahumara from Municipio of Guazapares who is writ- 
ing a series of booklets in Tarahumara and Spanish under the direc- 


tion of Mr. Burgess (cf. Mares, 1972 and 1974). 


(85 | 


Or 


have commented on the many uses and functions of 
maguey or the mescal plant. Ina future paper, Bye will 
review these reports with respect to the T'arahumara as 
2n whole. We now propose to review the utilization and 
significance of the magueys to one group, the Western 
Tarahumara‘’. This paper is the first part in a series on 
Western Tarahumara ethnobotany in which we plan to 
present information on and discussions of plants used 
by the natives living on the eastern slopes of the Rio 
Chinipas drainage area in the Municipios of Guazapares, 
Chinipas and Uruachi in southwestern Chihuahua. The 
Rio Chinipas, a tributary of the Rio Fuerte, originates 
west of the Continental Divide near Creel. 

Kight species of leave which occur in western ‘Tara- 
humara region will be discussed’. Most of the species are 
restricted to the northern Sierra Madre Occidental and 
range from the arid sub-tropical vegetation of the barran- 
cas to the cool pine forests of the sierras. Table I sum- 
marizes the range, elevation and ecological association 
of the Western Tarahumara agaves. 

UTILIZATION 

Agave is first and foremost a source of food. The com- 
mon method in preparing the heart (.e., the secondary 
thickened stem with compacted nodes) and leaf bases 
consists of pit-baking, usually by the men. Plants are 
selected by size and degree of development of the inflo- 
rescence bud, The leaves near the centre around the 
swollen flower stalk bud are usually disposed in a series 
of smaller leaves contrasting with the previous larger 
ones. Harvesting the plant Just prior to the elongation of 
the large inflorescence when the concentration of energy- 

'Tarahumara Baja, according to Mexico’s Instituto Nacional Indi- 
genista. 


” Consideration of other species found to the east and not recognized 
by the Western Tarahumara will be deferred to a later paper. 


if a] 
86 | 


rich sugars is highest is done by all people who eat ma- 
guey hearts. Pit-baked magueys are an important food 
source at the time of year when food stores of the pre- 
vious growing season are dwindling and before the arrival 
of summer rains which are needed for the growth of wild 
greens and cultivated plants. Occasionally, magueys are 
pit-baked at other times of the year. 

The plants are dug up with a pointed stick, an iron 
bar or an old ax. ‘Then the outer leaves are trimmed 
with a machete, leaving the hearts and leaf-bases. A 
wooden hook is inserted into the bottom of each heart, 
two of which are tied together and placed over a burro 
to carry to the pit. If people are carrying the hearts and 
the distance to the pit is great, the leaf-bases are cut off 
and carried separately in a basket. 

A fire, usually of green oak wood (Quercus spp.) is 
built in the pit with rocks placed on top of the wood. 
Hard, heavy rocks are chosen, since they hold heat longer. 
The round pit is usually about three feet deep and five 
feet wide. The same pit is used year after year. About 
noon, when the fire has burned down to coals and the 
smoke, which would give the hearts a bad taste, no longer 
comes out, the pit is ready for use. If the coals are too 
hot, a layer of earth is placed over them. Then the mes- 
‘al hearts with the leaf-bases still intact are placed in the 
pit. Sometimes a stick of ‘‘rolobusi’’® (Bouvardia sp.) 
is inserted into each heart to assure sweetness in cooking. 
If ‘‘rolobusi”* is not available, leaves of ‘‘usabi’’ (Prunus 
Gentry) are placed on the hot rocks at the bottom for 
the same purpose. After the hearts are in the pit, the 
pit is covered with leaves of a palm (Sabal wresana). 
Then a layer of soil is placed over the top and left for 

° Western Tarahumara belongs to the Uto-Aztecan family of lan- 
guages. It is written here using the Spanish orthography with the 


addition of the glottal (*). The r’s and 1’s differ from Spanish in that 
they are retroflexed. 


= 


[ 87 | 


two nights. On the morning after the second night, if 
the tops of the mescals are not done, the hearts are re- 
moved and the fire rebuilt under the rocks. Then the 
hearts are turned over, replaced in the pit, covered again 
and left for another day. If still not sufficiently cooked, 
they may remain in the pit-oven an extra day. 

Upon removal, the hearts and leat-bases are loaded 
into gunny sacks and baskets and taken to the house. 
They are either eaten by themselves or oftentimes with 
acorn drink, *‘gue huali’’ (Spanish: esquiate), or a corn 
gruel, ‘‘atonali’’ (Spanish: atole). The sweet flesh of the 
heart is eaten and the fibres spit out. The heart 1s some- 
times ground and mixed with tortilla dough. The prepa- 
ration and consumption of agaves have been described 
for other areas of the Tarahumara region (Pennington, 
1963: 129-131; Bennett and Zingg, 1985: 148-149). 
The hearts of Agave pacifica, A. Shreve, A. multifilifera 
and f. polianthiflora and the leaf-bases of 4. americana 
var. evpansa, A. multifilifera and A. polianthiflora are all 
pit-baked and eaten. 

‘Sugui’’ (Spanish: tesgiiino), a fermented beverage, 
‘an also be made from the well cooked mescal. In areas 
where agaves are plentiful, only the leaf-bases are used. 
In other areas, the hearts are also used. ‘The mescal is 
thoroughly mashed in a trough or hollow rock with a 
large mallet cut from the limb of an oak (Quercus spp. ). 
The juice is collected and strained, first through a coarse 
basket, then a cloth. After straining, the sweet Juice is 
cooked ina large pot by women, until all of the impuri- 
ties have come to the top and are skimmed off. Follow- 
ing the cooking, the juice is placed in a large-mouthed 
pot to cool and then put into a fermenting pot, ‘‘sicoli 
ronela’’ (Spanish: botija), where it is left for two or three 
days. The process can be speeded up by adding the juice 
of sprouted corn (Zea Mays) that has been ground, cooked 


. 


| 88 | 


and strained. Water is added if the ‘“‘imé”’ juice is too 
strong. When the bubbling of the fermenting process 
begins to slow down, a handful of ground wheat (7'riti- 
cum aestivum) is added. Then, within five to six hours, 
the ‘‘sugui™’ is ready to drink. If the drink is not to be 
used for several days, the pot is sealed with a board and 
mud. This will keep the drink from going bad for six 
to eight days. Agave bovicornuta is said to make a 
stronger ‘‘sugui’” than the other species. 4. americana 
and 4. vilmoriana are the only species not used for 
making the fermented drink. 

Tarahumaras also dry the roasted mescal so that it will 
last five to six months (during the dry season). The outer 
fibres are removed from the cooked leaf-bases and the 
meat is mashed, as when making “‘sugui’’, and then 
ground on the ‘‘majta’’ (Spanish: metate). It is then 
shaped into a cylindrical cheese form and placed in the sun 
to dry for a week. This *‘mesagoli”” or “‘imé cha’ poli’, 
mescal bread, is especially good food for long trips. It is 
‘carried in a bag of ‘‘cobisi’’ (Spanish: pinole), toasted 
corn grain that is ground dry into a powder and later 
mixed with water to drink, and used as a trail food. 


. 


al. pacifica is not used to make ‘‘mesagoli’’, since it is not 
so sweet, has less meat and the fibres are harder to chew. 

The emergent flower stalks, ‘‘balila’* (Spanish : quiote), 
are eaten as well. A young, tender stalk can be cut from 
the plant and roasted on coals. Then it is peeled and 
eaten. The taste is similar to that of the squash. It can 
also be chopped up, boiled, and mixed with the whey 
left over from making ranch-style cheese. When a stalk 
has elongated about a yard, the top foot is cut off and 
the remainder is peeled and roasted in the pit-oven along 
with mescal hearts and leaf-bases. The decapitated plant 
is allowed to grow and is cut the following year for pit- 
baking when it is sweeter. In other parts of the Tara- 


"89 | 


humara region, the prepared stalks are eaten with ‘‘gue’- 
huali’’ (Spanish: esquiate), toasted corn ground with 
water, and ‘‘cobisi’’, but this practice is not common in 
the western region. ‘The stalks of species of the subgenus 
Agave are eaten, while those of the members of sub- 
genus Littaeca are not because of their bitterness or 
small size. 

If the stalks be allowed to develop, the unopened flow- 
ers with the inferior ovaries removed are cooked and 
washed to remove the bitter constituent. The cooked 
flowers can be eaten boiled, fried or made into tortillas. 
The sweet nectar is likewise gathered with a small tube 
for consumption. The flowers of sfeave bovicornuta, A. 
pacifica, A. Shrevet and A. Wocomahi are eaten. The 
flowers of 4. bovicornuta are preferred as food because 
of their better taste and as they are available when there 
is little other food. On the other hand, flowers of 4. 
Wocomah, which appear during the rainy season, are not 
often collected, since there are many other preferred 
greens (Spanish: quelites) available at that time. 

Tortillas made from the perianth of the maguey flow- 
ers are especially valued by Western Tarahumara. The 
unopened flowers are cut from the large inflorescence. 
Usually a hooked pole is used to gather the flowers, so 
that the stalk is not broken, and the underdeveloped 
flower buds are allowed to develop tor future gathering. 
The flowers are put on the house roof and collected early 
the next morning, when they are much sweeter. They 
are cooked well and washed overnight in the arroyo to 
remove the bitterness. The next day they are ground 
with lime-cooked maize, or “*najpili” (Spanish: nixta- 
mal), and made into tortillas. Some cooked flowers are 
retained for later frying to be eaten with tortillas. 

Fibre is extracted from the leaves of Aeave multifih- 
fera and A. pacifica. The larger leaves are cut, beaten 


— 90 


and placed in water. The tissue surrounding the fibre 
bundles rots away. The resulting fibres are washed in 
the arroyo and commonly used in making lassos and 
domestic rope. 

Stupefying fish with agaves is well known among the 
Tarahumara (cf. Gentry, 1972: 11, 89). The Western 
Tarahumara use Agave Vilmoriniana, which is rich in 
sapogenins, and 4. bovicornuta and A. Wocomahi, the 
toxic constituents of which are unknown, to stupefy fish 
trapped in rock enclosures in the rivers. The plants are 
mashed on a rock, the juices are allowed to drain into 
the river and, finally, the mashed plants are thrown into 
the water. Several plants are needed rapidly to stupefy 
the trapped fish which, upon reaching the surface, are 
collected and prepared. 

The crushed leaves of Agave Vilmoriniana, which is 
rich in the sapogenin smilagenin, are prized as soap for 
bathing and washing clothes, wool, blankets and dishes. 
Not all ‘‘ahué”” leaves are good for washing. The whiter 
leaves are chosen over the greener ones which produce 
too strong a soap, especially for washing hair. Another 
method of selection is to twist the end of the leaf. If it 
breaks, that leaf is not used. 

Ritual curing with agaves is important to the Tara- 
humara ceremonies (Lumholtz, 1902: 821-822, 368). 
Small plants of fave Shreve: are placed in pots of 
water in front of the cross during fertility rites for sheep 
and goats and in ceremonies for the dead. 


Discussion 


About 300 years ago, the Tarahumara retreated west- 
ward and assimilated various tribes of close cultural and 
linguistic affiliations (Pennington, 1986: 8, 9, 229, 2380, 
Map I). Today they number about 50,000 (Pennington, 
1963: preface). The Western Tarahumara are geographi- 


91) 


‘ally isolated from the mainstream 'Tarahumara and live 
east of the Guarijio (Warihio). They number tewer than 
10,000 (Burgess, 1970: 64). 

The Western Tarahumara recognize two groups of 
Agave’. They recognize the group ‘‘imé™” (subgenus 
Agave) through its leaf with terminal and marginal 
spines and through the paniculate inflorescence. The 
‘imé”” group includes: *‘galime’’ (fl. americana var. ev- 
pansa), “sa pult”” (A. bovicornuta), ““gusime™” (4. paci- 
fica), “‘o'tosi™” (CL. Shrever), and *‘ojcome™” (P. Woco- 
mahi). The introduced succulent, -floe vera (Liliaceae), 


superficially resembling the maguey plant, is not widely 
considered an *‘imé™ although it is called *‘imé me’ ta- 
gochi’. The second group (subgenus Littaea) is recog- 
nized through its leaf with a terminal spine and smooth 
margin and its racemose inflorescence. This group in- 
cludes ‘‘chahut’’ (Ll. multifilifera), rv yéchili” (A. polian- 
thiflora) and “tahué™” (A. vidmoriana). The group carries 
no generic term. Some Western ‘Tarahumara include 
“chahul” and ‘ra’yéchili”” in the ‘‘imé*” group because 
they are pit-baked and eaten. 

There isa striking similarity and contrast in the recog- 
nition of the species of .feave among the Western ‘Tara- 
humara, the Guariyio to the west and the Tarahumara 
to the east. It may be explained by the geographical 
distribution of the magueys and, in part, by the histori- 
cal past. Table Ll summarizes the names of the eight 
species of -fgave occurring in the Western ‘Tarahumara 
region. 

In subgenus Littaca, Agave multifilifera and A. Vil- 
morimana are similarly recognized and domestically used 
by all three groups. One possible explanation is that the 

“The Spanish word, ‘“mescal’’, is locally used for the plants of 


Agave, but in other parts of Mexico this term is applied to a distilled 
beverage made from the Agave plant. 


leaves and fibres of ‘‘chahui’’ and the sapogenin-rich 
leaves of ‘‘ahué’” can be easily transported and traded 
throughout the area. On the other hand, food and fer- 
mented beverage made from the members of subgenus 
A gave are prepared and consumed near dwellings and 
‘an not easily be transported (except mescal bread). A. 
polianthiflora is presently known only from Western 
Tarahumara region and adjacent Sonora. fl. americana 
var. evpansa 1s not reported from the Guarijio or Tara- 
humara regions. 

Another significant point is that the species of eave 
here discussed are commoner on the western slopes and 
barrancas of the Sierra Madre Occidental. The diversity 
of species of subgenus f.2ave is greater toward the west 
(cf. Gentry, 1972). The increased diversity of ‘‘imé”’ 
encountered by the westward moving ancestors of West- 
ern Tarahumara could be handled by an expanded no- 
menclature borrowed from an assimilated tribe, such as 
the Guazapar. The language of the Guazapar was said 
to be the same as Guarijio, although it resembled Tara- 
humara (Pennington, 1968: 9). 


Boranical List or A gave 
Subgenus Agave 


Agave americana LL. var. expansa (Jacobi) Gentry, 
U.S.D.A., Agric. Handbook no. 899, p. 80. 1972. 
Western Tarahumara: GALIME (house maguey) 
Mexican: mescal maguey 
This large maguey, cultivated near houses, produces 
easily transplanted sucker shoots or ‘‘ranala’’ (Spanish: 
hijos). Its antiquity among the Western T'arahumara is 
unknown, and it is not reported from any other nearby 
regions of southwestern Chihuahua and adjacent Sonora. 
Gentry (1972: 84) suggests that it was introduced north- 


| [ 93 ] 


ward into Sonora and southwestern United States, proba- 
bly as an ornamental, after 1850. 

The bases of large leaves, pit-baked and eaten or pre- 
pared as mescal bread, are sweet, bland and lack fibres. 
One large plant usually provides sufficient material to 
fill a large baking-pit. Since other parts of the plant— 
the stalk, flowers and heart—are bitter, they are not 
consumed. 


Specimen: Curmmuanua: Municipio de Guazapares, Rocoroibo, ca. 
6000 ft. Cultivated plants about 6-8 feet high near governor’s house, 
associated with spineless Opuntia, apple and peach trees. Nov. 10, 
1973. Bye and Burgess 5827. 

Onservation (by R. Bye): 

Cuimuanvua: Municipios de Guazapares y Chinipas, Wasachi, ca. 
6000 ft. A recently transplanted linear plantation near walls of a 
ranchito in the open flat above arroyo, July 23, 1974. 


Agave bovicornuta Gentry, Carnegie Inst. Wash. 
Pub. 527, p. 92. 1942. 

Western Tarahumara: sA°’puLi (meat barranca: mean- 

Ing uncertain) 

Mexican: mescal lechuguilla 


This attractive maguey is found in the middle zone of 
the barrancas associated with oaks (Quercus spp.). The 
flowers, well washed to remove the bitterness, are eaten 
cooked or in the form of tortillas and are preferred to 
those of other species in making tortillas. The hearts are 


. 


baked and used in preparing a strong “‘sugui’’ which is 
said to be similar to ‘*pisto’’, the local Mexican distilled 
alcohol from cfeave and Yucca. The caustic juice of the 
leaves causes a burning irritation, followed by white blis- 
ters, on sensitive skin. The plant is employed to stupefy 
fish. 

Specimen: Cuiuuanua: Municipios de Guazapares y Chinipas, be- 
tween Cusdrare and Nopalero, ca. 3500 ft. Seattered plants on rocky 
slopes of the barranca associated with oaks (Quercus spp.) and below 
the mixed oak and pine forests. Nov. 8, 1973. Bye, Burgess and 
Mares 5806. 


94 | 


Opservation (by R. Bye): 

Curnuanua: Municipios de Guazapares y Chinipas, between Wasa- 
chi and La Paz, ca. 5000. Scattered plants in the lower section of 
the mixed oak and pine forest. The leaves are a light yellow-green. 
July 23, 1974. 


A gave pacifica 'Trel., U.S. Natl. Herb. Contrib. 28: 
118. 1920. 
Western Tarahumara: GUSIME (maguey of the woods 
or stick maguey) 
Mexican: mescal del monte (wild plant) 
mescal casero (cultivated plant) 


Found in the lower parts of the arid sub-tropical bar- 
rancas in short thorn forest, fave pacifica produces 
sucker shoots which are transplanted near dwellings in 
the lower oak (Quercus albocincta) zone of the middle 
barranca zone. Considered the most delicious and diffi- 
cult to collect in the field, this species is often grown 
near the houses. It also became popular with the 
Spaniards and Mexicans who settled in the deep can- 
yons and later transplanted and cultivated it. 

The hearts of the cultivated plants are larger than 
those of the wild plants. The baked hearts are sweet and 
consumed baked or in the form of fermented drink. The 
flowers are eaten cooked or in the form of tortillas. The 
fibres from the leaves provide cordage and thread. 

Specimen: Curmuanvua: Municipios de Guazapares y Chinipas, Cu- 
sirare, ca. 3000 ft. Cultivated plants, whose suckers originated from 
the wild in the lower parts of the barranca, in a hollow at the base 
of a field ‘‘trinchera’’ or stone retaining wall near a ranchito, Plants 
about 5 feet high with the ‘‘quiotes’’ cut off. Associated with such 
woody plants as /pomoea arborescens and Randia sp. Nov. 8, 19738. 
Bye, Burgess and Mares 5796. 

Opservation (by R. Bye): 

Curmuanua: Municipios de Guazapares y Chinipas, below La Paz, 
ea, 3500 ft. Cultivated plants along astone fence and margin of field 


near house surrounded by a peach orchard. In the lower oak (Quer- 
cus albocineta) zone. July 25, 1974. 


Agave Shreve: Gentry, Carnegie Inst. Wash. Pub. 
527, p. 95. 1942. 

Western Tarahumara: o'ros. (white, plural form) 

Mexican: mescal blanco 


This whitish appearing plant is scattered in open areas 
in the middle and upper zones of the barrancas in the 
oak and mixed oak and pine vegetation. The hearts are 
pit-baked for eating or are employed in making mescal 
bread or a fermented drink. The stalks are also cooked 
for food. Small plants are used in curing and in death 
ceremonies. 

Specimen: Curavuanvua: Municipios de Guazapares y Chinipas, No- 
palero, ca. 4000 ft. Small scattered plants on open, rocky slope with 
mixture of oaks (Quercus spp.), pines (Pinus spp.) and Nolina mata- 
pensis. Some plants with “‘quiotes’’ removed. Nov. 9, 19873. Bye 


and Burgess 5813. 


Agave Wocomaht Gentry, Carnegie Inst. Wash. Pub. 
527, p. 96. 1942. 

Western ‘Tarahumara: OJCOME (pine maguey) 

Mexican: mescal verde 


A small maguey, -feave Wocomahi is found scattered 
in rock crevices in the sierras among pines. Baked hearts 


‘an be used to make *‘sugui’’ and “‘mesagoli’’, although 


some hearts are said to have a bad taste and are not in- 
gested. The flowers are an acceptable food but are not 
eaten, as they appear at a time when there is an abun- 
dance of other greens. 

Specimens: Cuimuanua: Municipios de Guazapares y Chinipas, No- 
palero, ca. 4000 ft. Few scattered plants on open, rocky slope with 
a mixture of oaks (Quercus spp.) and pines (Pinus spp.). The plants 
are commoner above. Nov. 9, 1973. Bye and Burgess 5812.— Muni- 
cipio de Guazapares, Rocoroibo, ca. 6000 ft. On rocky, open slope 
above arroyo with mixed oaks (Quercus spp.), pines (Pinus Engelmannit 
and P. ponderosa) and madronos (Arbutus spp.). This plant was more 
robust than usual because of the moist site. Nov. 10, 1973. Bye and 
Burgess 58385. 


96 | 


Subgenus Littaea 


A gave multifilifera Gentry, U.S.D.A., Agric. Hand- 
book no. 899, p. 46. 1972. 
Western Tarahumara: CHAHUI 


A medium sized plant, ‘‘chahui’’ grows in rock out- 
crops in the upper zone of the barrancas in the mixed 
oak and pine vegetation. The hearts and leaf bases are 
pit-baked, eaten or made into ‘‘mesagoli’” and ‘‘sugui’’. 
The leaves are an important source of fibre. 

Specimens: Curnuanua: Municipios de Guazapares y Chinipas, near 
Nopalero, ca. 4000 ft. In crevices of rocks in the mixed oaks (Quer- 
cus spp.) and pines (Pinus spp.). Nov. 8, 1973. Bye, Burgess and 
Mares 5783. —Municipio de Guazapares, along ridge above Nopalero, 
cea, 5000 ft. Few scattered plants in rock crevices on outcrops in 
mixed oaks (Quercus spp.), pines (Pinus spp.) and Arctostaphylos pun- 
gens. Nov. 9, 1973. Bye and Burgess 5819. 

Agave polanthifiora Gentry, U.S.D.A., Agric. 

§ i 
Handbook no. 399, p. 51. 1972. 
Western ‘Tarahumara: RI YECHILI 


This small, attractive and infrequent maguey grows 
on partially open rocky slopes and outcrops in the mixed 
oak and pine forest in the upper zone of the barrancas. 
The hearts and leaf bases were formerly eaten as food 
and employed to make a fermented drink. 

Specimens: Cutuuanua: Municipios de Guazapares y Chinipas, above 
Nopalero, ca. 4500 ft. Few scattered plants in whiterock crevice in 
open spots with oaks (Quercens spp.), pines (Pinus spp.) and Arcto- 
staphylos pungens. *“‘Quiotes’’? about one foot high. Nov. 7, 1973. 
Bye, Burgess and Mares 5779.— Below Wasachi, ca. 5500 ft. Few 
scattered plants on open, moist rocky surface with mixed oaks (Quer- 
cus spp.) and pines (Pinus spp.). “‘Quiote’’ about one foot high with 
flowers rose. July 23, 1974. Bye, Burgess and Mundy 6414. 

Agave Vilmoriniana Berger, in Fedde Repert. Spec. 

Nov. Reg. Veg. 12: 508. 19138. 
Western Tarahumara: AHUE 
Mexican: amole 

[ 97 | 


These spider-like plants cling to the vertical rock walls 
of the barrancas from the lower arid sub-tropical region 
to the higher cooler region. The plant is not eaten. The 
leaves provide a soap for bathing and washing clothes, 
wool blankets and utensils, while the macerated plants 
are used as a fish poison. 

Specimen: Cuimuanua: Municipios de Guazapares y Chinipas, be- 
tween Nopalero and Cusidrare, ca. 3500 ft. Scattered plants found on 
protected vertical walls. Nov. 8, 1973. Bye, Burgess and Mares 5805. 

CONCLUSIONS 

Kight species of fave are recognized and used by the 
Western Tarahumara. ‘The members of subgenus -f cave, 
generically called “‘imé’’, are distinguished from the 
members of subgenus Littaca. The two groups are differ- 
entiated on gross morphology of leaves and inflores- 
cences. Some Western ‘Tarahumara include all of the 
magueys that are pit-baked under “‘imé”’. 

ach species of “‘imé™” has a name which reflects the 
habitat preference or a characteristic of the species. Al- 
though the Tarahumara to the east also recognize sub- 
genus -feave, ““mé’, as different from that of subgenus 
Littaea, the Western ‘Tarahumara recognize a greater 
diversity of species, and their names are similar to 
Guarijio names. In subgenus Littaca, two of the three 
species on the eastern slopes of the Rio Chinipas area 
have names in common with Guarijio names to the west 
and ‘Tarahumara names to the east. The third species, 
A. polianthiflora, has not been reported east of the 
Western Tarahumara region. 

There is a greater specific diversity of .feave on the 
western slopes of the Sierra Madre Occidental than on 
the eastern slopes. The similarity of the Western ‘Tara- 
humara names to those of the Guarijio for the members 
of subgenus .f@ave may be explained, in part, by the 
western movement of ancestors of the Tarahumara from 


[ 98 ] 


the eastern foothills and uplands to the western uplands 
and barrancas. Subsequent assimilation of former tribes 
of that region may have included additions to Tarahu- 
mara 4 gave nomenclature. The similar names for Af. 
multifilifera and A. Vilmoriniana may reflect the wide 
familiarity of the leaf, fibre and soap products of the 
plants. These plants and products would probably be 
encountered in travel, transport and trading. The other 
species provide food and beverage prepared and gener- 
ally eaten near dwellings and usually not traded or 
offered to strangers. 

‘Two species of f4gave produce sucker shoots, and the 
Western Tarahumara have used this characteristic to ad- 
vantage in their propagation. The antiquity of propaga- 
tion of 4. americana var. expansa is unknown, but its 
presence in northwestern Mexico is thought to be only 
a century old. ‘The much valued “f. pacifica has been 
propagated closer to the locality of its preparation and 
consumption and at higher elevations than its normal 
range. 


SPECIMENS AND PHOTOGRAPHS 


Voucher specimens were collected, prepared and iden- 
tified by R. Bye. Dr. H. S. Gentry was very helptul 
in instructing him in the preparation and identification 
of sterile and fertile plants and specimens. ‘The speci- 
mens are deposited in the Economic Botany Herbarium 
of Oakes Ames in the Botanical Museum, Harvard Uni- 
versity (KCON). Duplicates will be distributed to the 
Jniversidad Nacional Aut6noma de México (MENU), 
Gray Herbarium of Harvard University (GH), and the 
Herbarium of H. S. Gentry in Phoenix, Arizona. 

All photographs were taken by Don Burgess. 


TABLE I 


Western Tarahumara Agave 


Ecological 
Range Elevation Association}! 
subgenus Agave 
americana w Mexico to sw U.S. (100)-6000 ft. cult. (PO) 
var. expansa Jalisco to Ariz. & Calif. 
bovicornuta nw Sierra Madre Occidental 3000-6000 ft. Q-PO 
Son., Chih. & Sin. 
pacifica nw Mexico (0)-3500 ft. sT 
Son., Chih. & Sin. (cult.sT-O) 
Shrevet nw Mexico 3000-6000 ft. O-PO 
Son. & Chih. 
Wocomaht nw Sierra Madre Occidental 5000-7500 ft. © PO-P 
Son., Chih., Sin. & Dgo. 
subgenus Litlaea 
multifilifera nw Sierra Madre Occidental 5000-6000 ft. O-PO 
Chih. 
polianthiflora nw Sierra Madre Occidental 4000-6000 ft. ©Q-PO 
Son. & Chih. 
Vilmoriniana w Mexico 2000-4000 ft. sT 


Son. & Chih. to Jalisco & 
Aguascalientes 


1—_cult.=cultivated; O=oak forest (Quercus spp., e.g. Y. albocincta); P=pine 
forest (Pinus spp., e.g. P. Engelmannii, P. arizonica, P. ponderosa); PO= 
pine-oak forest; sT=arid sub-tropical forest with Ipomoea arborescens, Bur- 


sera spp. and various woody Leguminosae. 


' 100 | 


TABLE II 
Nomenclature of Western Tarahumara Agave 


Species Guarijio! W. Tar. Tarahumara? 
subgenus Agave _ imé mé 
americana 0 galime 0° 


var. expansa 


borvicornuta —sapuri sa’puli _ 
pacifica — gusime ku’uri 
Shrevei totosa o’tosa mé* 
Wocomaht wocomahi ojcome mésagori 
mé 
subgenus Littaea 
multifilifera chahui chahui chawi (or chahui) 


polianthiflora taiehcholi ri’yéchili 0° 


Vilmoriniana hauwé ahueé awé (or ahué) 


1—Guarijio (or Warihio) names, from Gentry (1942 and 1963). 


2—Tarahumara names collected by R. Bye in the region of Creel-Barranca 
del Cobre-Barranca de Batopilas. 

3—Western Tarahumara names, plant photographs and specimens not recog- 
nized by native informants. 


4—One informant, who grew up west of Cerocahui near Western Tarahumara 
Zo 


region but now lives in Barranca de Batopilas, recalled “retosa” as the 
name used for this maguey in his youth. 
“0” =species not recorded from the area. 


“__” — data uncertain. 


( 101 | 


ACKNOWLEDGMENTS 


We would like gratefully to acknowledge the help and 
encouragement of our families and friends in the United 
States and Mexico who generously supported our studies. 
A special thanks is extended to those Tarahumara who 
through friendship shared their knowledge. Without the 
continual encouragement and critical evaluation of Pro- 
fessor C. W. Pennington, Department of Geography, 
Texas A & M University, the research upon which these 
notes are based might have faltered. Professor Richard 
Evans Schultes, Botanical Museum, and Dr. Bernice 
Schubert, Arnold Arboretum, kindly read the manu- 
script and offered suggestions. Robert Bye would like 
personally to thank Dr. Margaret A. Towle, Curator of 
the Ethnobotanical Laboratory of the Botanical Mu- 
seum, for her support and valuable suggestions. Financial 
support between 1971 and 1974 was generously extended 
to Robert Bye by the Botanical Museum and the De- 
partment of Biology, Harvard University, the National 
Geographic Society and the National Science Founda- 
tion Grant GB-35047 for Improvement of Doctoral 
Dissertation Research. 


[ 102 ] 


BIBLIOGRAPHY 


Bennett, W.C., and R.M. Zingg. 1935. The Tarahumara. Univ. of 
Chicago Press, Chicago. 


Burgess, D.H. 1970. ‘‘Tarahumara Phonology (Rocoroibo Dialect). ”’ 
In R.W. Ewton, Jr., and J. Ornstein (eds.). 1970. Studies in 
Language and Linguistics (1969-1970), pp. 45-66, Texas Western 
Press, El Paso. 


Castetter, E.F., W.H. Bell and A.R. Grove. 1938. ‘‘The early uti- 
lization and the distribution of Agave in the American Southwest. ”’ 
Univ. New Mex. Bull., Bio. Ser. 5(4). 


Gentry, H.S. 1942. Rio Mayo Plants. Carnegie Institution of Wash- 
ington, Publ. no. 527. 


——, 1963. ‘‘The Warihio Indians of Sonora-Chihuahua: an ethno- 
graphic survey.”’’ Bur. Amer. Ethnol., Bull. 186: 61-144, pl. 28-38, 


——, 1972. The Agave Family in Sonora. U.S.D.A., Agric. Hand- 
book, no. 399. 


Lumholtz, Carl. 1902. Unknown Mexico. Charles Scribner’s Sons, 
New York. Volume I. 


Mares Trias, A. 1972. Hue’cd E’cartigame Nehualiame Ju Imé. (Haec- 
mos muchas cosas con el mezcal). Instituto Lingiiistico de Verano 
en coordinacion con la Secretaria de Educacién Ptiblica de México, 
México, D.F. 


——, 1974, Rega Me’liame Ju Ye Ajagame. (Como cazamos y pes- 
camos). Instituto Lingiistico de Verano en coordinacioén con la 
Secretaraia de Educaci6n Publica de México, México, D.F. 


Pennington, C.W. 1963. The Tarahumar of Mexico. Univ. of Utah 
Press, Salt Lake City. 


[ 103 ] 


[ILLUSTRATIONS 


{ 105 | 


PLATE XVII 


7 = 
= 


PO, ON a: 
fae. cata 


Agave americana var, expansa (— galime’”) 


Mature leaves about 4) feet long. 


| 106 | 


PLaTE XVIII 


; 66 739 
Agave bovicornuta ( ‘sa’ puli’’) 


Mature leaves about 2 feet long. 


PLATE XIX 


ee 29) 


Agave pacifica (~ gusime 


Mature leaves about 3 feet long. 


[ 108 ] 


PLaTrE XX 


Agave Shrevei (“‘o Ai 


Mature leaves about 13 . feet long. 


| 109 | 


PLATE XX] 


Agave Wocomahi (‘ojcome’’) 


Mature leaves about | foot long. 


[ 110 | 


PLATE XXII 


hili’’) 


éc 
aves about 4 inches long. 


ri’yé 


e polianthiflora (** 


) 


4 gar 


N 


Agave multifilifera (“‘chahui’’) 


{ature le 


+ feet long. 


2 


Mature leaves about 


PLATE XNIII 


Agave Vilmoriniana (- ahué’’) 


Mature leaves about 5 feet long. 


[112 ] 


HARVARD UNIVERSITY 


CampripGre, Massacnuserts, Ocroser 31, 1975 VoL. 24, No. 6 


NUTRITIONAL VALUE OF COCA 
BY 
JAMES A. Duke’, Davip AULIK* AND 
Timoruy PLlowMan® 


Leaves of wond’rous nourishment 

Whose Juice Suce’d in, and to the Stomach tak’n 

Long Hunger and long Labour can sustain; 

From which our faint and weary Bodies find 

More Succor, more they cheer the drooping Mind, 

Than can your Bacchus and your Ceres join’d. 
—AsBrAHAM Cow ry in Mortimer’s 

History of Coca 


Abstract. Coca leaves (Erythroxvylum Coca am.) 
from Chapare, Bolivia, compared to an average of 50 
other Latin American vegetable products, are higher in 
calories (805 per 100 g compared to 279), protein (18.9 
g: 11.4 g¢), carbohydrate (46.2 g: 87.1 g), fiber (14.4 g: 
3.2 g), ash (9.0 g: 2.0 g), calcium (1540 mg: 99 mg), 
phosphorus (911 mg: 270 mg), iron (45.8 mg: 3.6 mg), 
vitamin A (11,000 IU: 185 LU), and riboflavin (1.91 
mg: 0.18 mg). Coca was lower than the average for the 
50 plant foods in oil content (5.0 g per 100 g compared 
to 9.9 g), moisture (6.5 g: 40.0 g), thiamin (0.35 mg: 
0.388 mg), niacin (1.8 mg: 2.2 mg), and ascorbic acid 

‘Chief, Plant Taxonomy Laboratory, Plant Genetics and Germ- 
plasm, Institute, Agricultural Research Service, Beltsville, Maryland. 

> WARE Institute, Inc., Box 2599, Madison, Wisconsin. 


3 Fy ° 7 F . 
Research Associate in Economic Botany, Botanical Museum, 
Harvard University, Cambridge, Massachusetts. 


[ 113 ] 


(1.4 mg: 18.0 mg). Ingestion of 100 g of the Bolivian 
coca leaves tested would more than satisfy the Recom- 
mended Dietary Allowance for reference man and 
woman of calcium, iron, phosphorus, vitamin A, vita- 
min B, and vitamin EK. However, the leaves also contain 
alkaloids and may harbor pesticide residues. 

Unable to establish the nutritional value of coca leaves 
(Mrythrovylum Coca Lam. )afterconsulting many sources, 
we obtained a one kilogram sun-dried sample from San 
Francisco, Province of Chapare, Bolivia, in June, 1974. 
Using methods listed in References and Notes, we ob- 
tained the following nutritional analysis (1): calories, 805 
per 100 @: moisture, 6.5 g; protein, 18.9 @: carbohy- 
drate, 46.2 @; fat, 5.0 g: vitamin A, 11,000 LU (as 
beta-carotene); vitamin C, 1.4 mg; vitamin B, (thia- 
mine), 0.85 mg; vitamin B, (riboflavin), 1.9 m@: niacin, 
1.29 meg: calcium, 1,540 mg: iron, 45.8 me: vitamin 
Ki, 43.5 [LU (as d-alpha tocopherol): vitamin B,, 0.508 
mg: folicacid, 0.1830 mg: vitamin B,,, 1.05 meg: iodine, 
5.0 meg: phosphorus, 911 mg: magnesium, 213 mg; 
zinc, 2.70 mg; copper, 1.21 mg: biotin, 0.0868 mg; 
pantothenic acid, 0.684 mg: and sodium, 40.6 mg. The 
analysis of other elements by emission spectroscopy 
yielded the following amounts: potassium, 2.02 @ per 
r: stronti- 


S 


_ 


100 @; aluminum, 89.5 mg; barium, 4.67 m 
um, 9.71 mg; boron, 5.85 mg: zine, 2.70 mg; manga- 
nese, 6.65 mg: and chromium, 0.359 mg. 

Surprised by the high values, especially in calcium and 
iron, we tabulated nutritional averages for other plant 
products ingested by Latin Americans (‘Table 1). Com- 
pared with an average from ten nuts and oilseeds (2) 
(Sesamum indicum, Terminalia Catappa, Prunus A myg- 
dalus, Corylus spp., Arachis hypogaca, Castanea spp., 
Bertholletia evcelsa, Helianthus annuus, Anacardium 
occidentale and Inga spp.), the San Francisco coca leaves 


( 114 ] 


were higher in protein, carbohydrate, ash, calcium, phos- 
phorus, iron, vitamin A and riboflavin. Coca was lower 
in calories, moisture, fat, thiamin, niacin, and vitamin 
C. Compared with an average of ten pulses (Vigna un- 
guiculata, Cicer arietinum, Cajanus Cajyan, Pisum sativum, 
Vicia Faba, Phaseolus vulgaris, Dolchos Lablab, Lens 
spp., Glycine Max and Lupinus mutabilis), coca was 
equal in fat; higher in fiber, ash, calcium, phosphorus, 
iron, vitamin A, and riboflavin; and lower in calories, 
moisture, protein, carbohydrate, thiamin, niacin, and 
vitamin C. Compared with an average of ten cereals 
(Amaranthus caudatus, Oryza sativa, Avena sativa, Chen- 
opodium pallidicaule, Chenopodium Quinoa, Hordeum 
vulgare, Secale cereale, Coix Lachryma-jobi, Zea Mays 
and Triticum aestivum), coca was higher in protein, fat, 
fiber, ash, calcium, phosphorus, iron, vitamin A, ribo- 
flavin, and vitamin C; lower in calories, moisture, carbo- 
hydrate, thiamin, and niacin. Compared with an average 
of ten vegetables (Canna edulis, Capsicum spp., Allium 
sativum, Arracacha wanthorrhiza, Ipomoea Batatas, Cy- 
clanthera pedata, Cucurbita mavima, Allium Cepa, Bras- 
sica oleracea, and Tropaeolum tuberosum) and an average 
of ten fruits (Persea americana, Ananas comosus, Musa 
sapientum, Cocos nucifera, Passiflora mollissima, Annona 
Cherimolia, Prunus persica, Fragaria spp., Annona mu- 
ricata, and Meus Carica), coca was high on all counts 
except moisture and vitamin C. 

The present coca analyses are comparable to an average 
of three earlier coca analyses from Bolivia (8) and three 
recently reported from Peru (4). Frequent reports that 
coca has no nutritional value should be re-evaluated in 
view of these findings. The comparatively high nutri- 
tional values for coca are due partly to the fact that the 
leaves are dry (less than 10% moisture) when purchased, 
whereas most other foods are higher in moisture. 


[ 115 | 


In most areas where it is used, coca should be con- 
sidered a masticatory since it is not wholly consumed by 
the chewer. Typically, the leaves are first moistened in 
the mouth with saliva, then formed into a quid with the 
tongue and pushed into the upper cheek cavity. They 
are then sucked to extract the rich, green Juice which is 
subsequently swallowed. Usually some form of alkali 
is added to facilitate this extraction. When the chew is 
exhausted, it is usually spat out. Thus, the full comple- 
ment of nutrients present in the coca leaf is not consumed 
entirely, and the nutritional amounts reported here may 
be somewhat higher than the amounts actually ingested 
by the coca chewer. To our knowledge, no studies have 
been made on the nutritional value of the swallowed 
extract. 

In the Colombian Amazon, a variation of coca use is 
practiced by several tribes. Coca leaves are pulverized 
to a fine powder along with the ashes of Pourouma or 
Cecropia leaves. The mixture is placed in the mouth on 
the gums and inner cheeks and is eventually swallowed 
(8). In this case, and in instances where a coca chewer 
swallows his quid, the full complement of the leaf nutri- 
ents would be ingested. There is essentially no difference 
here between the use of coca and the direct consumption 
of food, in terms of nutrition. 

The amounts of coca consumed may contribute signifi- 
‘cantly to the diets of Andean coca chewers (5). If the 
average chewer ingests 60 g of Peruvian coca per day 
(5,6), he more than satisfies his requirements for calcium, 
even without the supplemental ash or lime usually added 
to the coca quid (6). No other food in the INC AP Food 
Composition Tables (2) approaches coca for calcium con- 
tent (1,789 mg). Other food items are high in calcium: 
sesame seeds, 1212 mg per 100 @: spinach flour, 488 mg: 
leaves of Laurus nobilis, 803 mg: leaves of Justicia pecto- 


[ 116 ] 


ralis, 668 mg; West Indian Almond ( Terminalia Catap- 
pa), 497 mg; powdered skim milk, 1,801 mg; whole 
milk, 921 mg; and alligator meat, 1,281 mg; but none 
equals coca. Few food plants can satisfy the calcium and 
iron in the Recommended Dietary Allowance (RDA) 
of reference man ingesting 100 g. The Bolivian coca 
leaves reported here do satisfy the RDA. 

Coca leaves may, however, contain 0.25 to 2.25% toxic 
alkaloids, including benzoylecgonine, benzoyltropine, 
cinnamylcocaine, cocaine, cuscohygrine, dihydroxytro- 
pane, hygrine, hygroline, methyleocaine, methylecgoni- 
dine, nicotine, tropacocaine, and a - and £ -truxilline 
(4,7). These alone could make the nutritious coca leaf 
undesirable as a source of nutrients. he average coca 
chewer could also ingest 442 mg of copper in a year if 
the San Francisco leaves are typical; but this amount 
is not excessive. 

Many coca growers in both high (Chapare, Bolivia) 
and low (Yungas, Bolivia) rainfall areas may use insecti- 
cides. The leaves are not intentionally washed, and har- 
vest and curing are timed to avoid rainfall. In considering 
coca for human consumption, the leaves should be ana- 
lyzed for insecticide residues. 

Although coca leaves contain relatively high levels of 
certain nutrients, the presence of alkaloids and the pos- 
sible presence of insecticide residues suggest caution in 
coca chewing. 


[ 117 ] 


REFERENCES AND NOTES 


Methods: Protein (Yo N 6.25), Association of Official Analyti- 
cal Chemists (A.O.A.C.), 11th Ed., 16. 1970; moisture, vacuum 
oven, A.O.A.C., 122. 1970; ash, A.O.A.C., 123. 1970; fat, 
A.O.A.C., 129. 1970; carbohydrates, by difference; calories, by 
calculation; vitamin A, Moore & Ely, Ind. Eng. Chem. Anal. 
Ed., 13: 600. 1941; vitamin C, J. Biol. Chem., 147: 399. 1948; 
vitamin B,, A.O.A.C., 771. 1970; vitamin Bz, A.O.A.C., 789. 
1970; niacin, A.O.A.C., 787. 1970; elemental, J.A.O.A.C., 51: 
10038. 1968; vitamin E, Acta Chemica Scandinavica 11: 34-43. 
1957; vitamin Be (Streptococcus carlsbergensis), Atkins, Schultz, 
Williams & Frey, Ind. & Eng. Chem., Anal. Ed., 15: 141. 1948; 
folic acid, A.O.A.C., 786. 1970; vitamin B,2, U.S.P. 17: 864. 
1965; iodine, ashing, A.O.A.C., 674. 1970; colorimetry, W.T. 
Binnerts, Anal. Chemica Acta 10: 78. 1954: biotin (Lactobacillus 
arabinosus), Wright & Skeggs, Proc. Soc. Exp. Biol. & Med., 56: 
95, 1944; pantothenic acid, Nielands & Strong, Arch. Biol., 19: 
2. 1948, 


Wu Leung, W.& M. Flores. 1961. Tabla de Composicién de Ali- 
mentos para Uso en América Latina. Instituto de Nutrici6n de 
Centro América y Panama (INCAP) and Interdepartmental Com- 
mittee on Nutrition for National Defense (ICNND). U.S. Govern- 
ment Printing Office. Washington, D.C. 


del Granado, J.T. 1931. Plantas Bolivianas. Arno Hermanos. 
La Paz. 
Machado, E. 1972. El género Erythroxylon en el Peru. Ray- 


mondiana 5: 5-101. 


Hanna, J.M. 1974. Coca Leaf Use in Southern Peru: Some Bio- 
social Aspects. Am. Anthropologist 76 (2): 281-296. 


Baker, P.T. & R.B. Mazess. 19638. Calcium: Unusual Sources in 
the Highland Peruvian Diet. Science 142: 1466-7. 


Willaman, J.J. & B.G. Schubert. 1961. Alkaloid-bearing plants 
and their Contained Alkaloids. U.S. Department of Agriculture. 
Washington, D.C. 


Schultes, R.E. 1957. A new method of coca preparation in the 
Colombian Amazon. Bot. Mus. Leafl. Harvard Univ. 17 (9): 241- 


246, 


[ 118 ] 


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[119 J 


a4 


DE PLANTIS TOXICARIIS EF MUNDO 
NOVO TROPICALE COMMENTATIONES XIII 


NOTES ON POISONOUS OR MEDICINAL MALPIGHIACEOUS 
SPECIES OF THE AMAZON 
BY 
Ricuarp Evans ScCHULTES 


This paper is offered in continuation of a series pre- 
senting ethnopharmacological observations made in the 
northwest Amazon. The malpighiaceous species enumer- 
ated in the following pages—all apparently unstudied 
from the phytochemical and pharmacological points of 
view—have been found to have interesting applications 
amongst the Indians of the region. 

The identifications of the voucher specimens were 
made by Dr. José Cuatrecasas of the Smithsonian In- 
stitution and are cited and described in his monograph 
of Colombian Malpighiaceae which appeared in Webbia 
13 (1958) 843-664. The drawings herein produced were 
prepared by Mr. Joshua B. Clark. 

The ethnopharmacological observations were made by 
the writer in the Amazonian basin of Colombia between 
1941 and 1954. They are offered in the hope of stimu- 
lating research to discover new biodynamic principles or 
to find new potentially valuable medicinal plants. Re- 
search along these lines is especially needed for this family 
of 60 genera and more than 880 species which has re- 
cently been described as ‘‘scarcely touched, chemotaxo- 


[121 J 


nomically speaking’. (Gibbs, R.D. ‘‘Chemotaxonomy 
of Flowering Plants” 8 (1974) 1676-1677.) 

The best known members, from the point of view of 
their phytochemistry, are undoubtedly the several closely 
allied species of Banisteriopsis—B. Caapi, B. inebrians, 
B. Rusbyana—employed in northern South America in 
elaborating the narcotic drink known variously as aya- 
huasca, caapi, natema, pinde and yajé. ‘These species 
contain psychoactive alkaloids—the first two, B-carboline 
alkaloids; thethird, N, N-dimethyltry ptamine( Deulofeu, 
V. ‘Chemical Compounds Isolated from Banisteriopsis 
and Relative Species’ in [| Ed.D. Efron] ‘‘Ethnopharm- 
acologic Search for Psychoactive Drugs’” Public Health 
Service Publ. No. 1645, Washington, D.C. (1967) 893- 
402). 

One hundred and twenty-five years ago, when Spruce 
collected the type material of Banisteriopsis Caapi and 
described its narcotic use in the northwest Amazon of 
Brazil, he despatched material for chemical analysis— 
material which was not analyzed until 1968 (Schultes, 
R.E., B. Holmstedt and J.-E. Lindgren *‘De Plantis 
‘Toxicariis e Mundo Novo Tropicale I11. Phytochemical 
Kxamination of Spruce’s Original Collection of Banis- 
teriopsis Caapr” in Bot. Mus. Leafl. Harvard Univ. 22 
(1969) 121-182). Spruce was struck by the presence in 
this family of such a potent narcotic. In 1852, he wrote: 
‘2... Lsaw, not without surprise, that it belonged to 
the order Malpighiaceae and the genus Banisteria, of 
which I made it out to be an undescribed species. .. . 
My surprise arose from the fact that there was no narcotic 
Malpighiad on record, nor indeed any species of that 
order with strong medicinal properties of any kind... . 
The seed [of Bunchosia] is described in books as poison- 
ous, and if it be really so, then it is the only instance, 
so faras I know, of the existence of any hurtful principle 


[ 122 ] 


in the entire family of Malpighiads, always excepting 
that of the Caapi. Yet strong poisons may lurk undis- 
covered in many others of the order, which is very large 
.... (Spruce, R. [Ed. A.R. Wallace] ‘‘Notes of a 
Botanist on the Amazon and Andes”’ 2 (1908) 421-422). 


Banisteriopsis Martiniana (Juss.) Cuatrecasas 
var. laevis Cuatrecasas in Webbia 18 (1958) 502. 

Cotompia: Comisaria del Amazonas, Vaupés, Rio Apaporis, Raudal 
de Jirijirimo. H. Garcia-Barriga 13706.—Same locality. November 
25,1951. R.E. Schultes et I, Cabrera 14569, —Comisaria del Amazonas, 
Rio Apaporis, Soratama. August 20, 1951. Schultes et Cabrera 13615. 


Garcia- Barriga (Flora Medicinal de Colombia 2 (1975) 
69) has recently reported that Banisteriopsis Martiniana 
var. /aevis may be used by the Makuna Indians of the 
middle Apaporis as one source of the narcotic prepa- 
ration yajé. 


Heteropteris riparia Cuatrecasas in Webbia 13 
(1958) 483. 


Cotomsia : Comisaria del Putumayo, Rio Putumayo, Puerto Ospina. 
Alt. 300 m. ‘‘Extensive liana along river’s edge.’’ July 1942. R.E. 
Schultes 4029.—Comisaraia del Vaupés, Rio Apaporis, Raudal de 
Jirijirimo. Alt. 250 m. June 12, 1951. R.E. Schultes et I. Cabrera 
12428,—Comisaria de] Amazonas, Rio Apaporis, Soratama. June 18, 
1951. Schultes et Cabrera 12654.—Comisaria del Vaupés, Rio Apaporis, 
Jinogojé (near mouth of Rio Piraparana). Alt. 260 m. ‘Vine. Flowers 
yellow.’’ June 5, 1952. Schultes et Cabrera 16591.—Same locality. 
June 8, 1952. Schultes et Cabrera 16665. 


The bark of this extensive liana is reported in these 
widely separated localities to be extremely poisonous, 
but the Indians in the middle Apaporis rasp the bark 
and prepare a tea which is employed internally in the 
treatment of gonorrhoea. The Makuna name on the Rio 
Apaporis 1s mee-see-gaw. 

Saponins, tannins and phenolic acids have been re- 
ported from the genus Heteropteris (Gibbs, loc. cit. ). 


| 223. 


Heteropteris macrostachya 4. Jussieu Malpigh. 
Synop. (1840) 275. 

CoLtombBia: Comisaria del Amazonas, Rio Apaporis, Soratama. Alt. 
250 m. ‘‘Bejuco; hojas doradas envés; flores amarillas. Nombre en 
lengua ‘geral’ capituriiva.’’ August 16, 1951. R. FE. Schultes et I. 
Cabrera 13533, 

The seeds of Heteropteris macrostachya are reputedly 
taken inatea by the Taiwano Indians of the Rio Kana- 
nari to treat diarrhoea. 

It may be significant that the seeds of Heteropteris 
suberosa Griseb., commonly called sarabatuciv, are simi- 
larly employed on the Rio Mauhés in the central Amazon 
of Brazil (Le Cointe, P.: **A Amazonia Brasileira’ 3 
(1934) 406). 

A collection of Hleteropteris macrostachya from Pana- 
ma (1..M. Johnston 1577) bears an annotation that the 
plant is ‘‘toxic™. 


Hiraea apaporiensis (watrecasas Webbia 18 (1958) 
404. 


Cotomsia: Comisaria del Vaupés, Rio Apaporis, Raudal Yayacopi 
(La Playa). August 18, 1952. R.E. Schultes et I. Cabrera 16969.— 
Jinogojé (near mouth of Rio Piraparana). August 25, 1952. Schulltes 
et Cabrera 17022,.—Comisaria del Vaupés, Rio Piraparana. August 
1952. H. Garcia Barriga 14215. 

The Maku Indians, who know Hiraea apaporiensis as 
ye-aing, employ a tea prepared from the leaves which is 
valued in the treatment of conjunctivitis. 


Hiraea Schultesii Cuatrecasas Webbia 18 (1958) 
408. 

CoLomspia: Comisaria del Vaupés, Rio Apaporis, Raudal Yayacopi 
(La Playa). March 16, 1952. R.E. Schultes et I. Cabrera 15996. 

A wash of the leaves of Hiraea Schultesii is valued by 
the Makuna Indians of the middle Apaporis in treating 
severe conjunctivitis. 


[ 124 | 


PiatTe AXTV 


paria 
Cuatr. 


pry 


€ 5 
(th. 


=—— 


So pe == 


Mascagnia glandulifera Cuatrecasas Webbia 13 
(1958) 865. 

Cotompia: Comisaria del Amazonas, Rio Apaporis, Soratama. July 
31, 1951. RE. Schultes et I. Cabrera 15208,—Same locality. August 
16, 1951. Schultes et Cabrera 13594. 

A poultice of crushed and boiled leaves of Mascagnia 
glandulifera is commonly applied to boils and similar 
infections by Indians of the middle Apaporis. 

Saponins have been reported from a species of this 
genus (Gibbs loc. cit. ). 


Mezia includens (Benth.) Cuatrecasas in Webbia 
13 (1958) 450. 

Cotompia: Comisaria del Vaupés, Rio Apaporis, Jinogojé (at mouth 
of Rio Piraparand) and vicinity. “‘Vine. Flowers bright yellow.” 
RE. Schultes et I. Cabrera 15691. 

This vine represents one of the medicinal plants with 
most uses amongst the Makuna Indians of the middle 
Apaporis. It is unusual that this should be so, in view 
of the relative scarcity of the plant. 

The bark, which contains apparently high concentra- 
tions of tannin, is prepared in an infusion which is taken 
warm in large quantities to treat urinary troubles and to 
provoke urination. Perhaps because of its diuretic proper- 
ties, it isemployed in the form of a tea in cases of swol- 
len legs—obviously edemas due to age and circulatory 
ailments. 

The root is considered a strong laxative: it is crushed 
and soaked together with water in which /arvva (the flour 
prepared from Manihot esculenta has been setting for 
several hours. 

The leaves, boiled for long periods into a tea, provide 
a strong emetic drink. They are also considered, when 
applied over the abdomen in the form of a cataplasm, a 
help in the treatment of what appears to be jaundice. 

The Makuna Indians call this vine ce’-taw-gai. 


[ 126 } 


PLATE XXV 


HIRAEA apaporiensis  cuatr. 


<a be . hy \ 
AIG \ yy 


WN 


ns 
Pé 


4 


\\ . \\ \ 
Sy \ \ 
NY 


Tetrapteris mucronata Cavanilles Diss. 9 (1790) 
4354, t. 262. 

CotomBia: Comisaria del Amazonas, Rio Apaporis, Soratama. June 
15, 1951. RE. Schultes et I. Cabrera 12554,—Comisaria del Vaupés, 
Rio Apaporis, mouth of Rio Pacoa. June 15, 1951. Schultes et Cabrera 
13559.—Rio Apaporis, Jinogojé (near mouth of Rio Piraparana). June 
20, 1952. Schultes et Cabrera 16771. 

The natives of the lower Rio Piraparana prepare a 
weak type of curare by boiling together for four or five 
hours the bark of Tetrapteris mucronata and of Strychnos 
Erichsonti R. Schomb. (Schultes et Cabrera 16770). 

Tetrapteris mucronata is said to be ‘employed by the 
Karaparana tribe in preparing yajé’ (Schultes et Cabrera 
12107). In this connection, it is of interest to note the 
report of the narcotic use of a species of Tetrapteris, 7. 
methystica (Schultes et Lopez 10184), in the Rio Negro 
basin of Brazil (Schultes, R. E. in Bot. Mus. Leafl., 
Harvard Univ. 16 (1954) 202-205. 


Tetrapteris silvatica Cuatrecasas Webbia 13 
(1958) 425. 

Cotompia: Comisaria del Vaupés, Rio Apaporis, Raudal Yayacopi 
(La Playa). August 18, 1952. R.E. Schultes et I. Cabrera 16960, 


The leaves of Tetrapteris silvatica are burned by the 
Makuna Indians, and the ashes, mixed with any oil or 
grease, are applied to what appear to be fungal patches 
on the skin. The Makunas refer to this vine as ¢fee-mee- 
a-mee-see-ma and recognize its very close relationship 
with the following species, Tetrapteris styloptera. 


Tetrapteris styloptera Jussieu in Ann. Sci. Nat., 
ser. 2 Bot. 13 (1840) 202. 

Cotompia: Comisaria del Amazonas, Rio Igaraparana, La Chorrera 
and vicinity. June 1942. R.E. Schultes 3914.—Same locality and 
date. Schultes 3927.—Rio Apaporis, Soratama and vicinity. August 


3, 1951. RIE. Schultes et I. Cabrera 12880,.—Same locality. August 


(128 ] 


Piatt XX VI 


MASCAGNIA) glandulifera  cuatr. 


[ 129 ] 


16, 1951. Schultes et Cabrera 13573, —Comisaria del Vaupés, Rio Apa- 
poris, mouth of Rio Pacoa. June 17, 1951. Schultes et Cabrera 12597 .— 
Jinogojé (near mouth of Rio Piraparana). June 5, 1952. Schultes et 
Cabrera 16591.—Same locality. June 8, 1952. Schultes et Cabrera 
16667.—Same locality. June 20, 1952. Schultes et Cabrera 16778.— 
Same locality. September 20, 1952. Schultes et Cabrera 17600. — Raudal 
Yayacopi (La Playa). August 18, 1952. Schultes et Cabrera 169523 ,.— 
Same locality and date. Schultes et Cabrera 16961.—Cachivera Jirijiri- 
ma. November 1951. H. Garcia-Barriga 13700, 


The ‘Tanimuka Indians on the Rio Miritiparana know 
this yellow-flowered vine as wee-po-awh. The bark is 
rasped and boiled to prepare an extremely bitter drink 
taken as a febrifuge. 

Amongst the Makunas, the leaves, reduced to ashes, 
are applied to itching infections of the skin that may be 
due to fungal growth. The ashes are mixed with oil or 
fat for application. This vine is one of the most impor- 
tant medicines amongst the Makunas, who have three 
distinct names for it: bce-ra-ree-a-ma, ho-ree-a-mee-see 
and né-na’-mee-hoo-ma. 


(130 | 


Puate XXVII 


TETRAPTERIS 
styloptera Juss. 


vA TETRAPTERIS 
| mucronata Cav. 


[131 | 


ECONOMIC BOTANY LIBRARY 
OF OAKES AME) 


RVARD BOTANICAL MUSEUM 


BOTANICAL MUSEUM LEAFLETS 
HARVARD UNIVERSITY 


CampripGr, Massacuusetrts, December 12, 1975 VoL. 24, No. 7 


STUDIES IN THE GENUS ONCIDIUM. IL. 
BY 
Joun E. Stacy* 


The genus Oncidium, one of the largest orchid genera 
in the New World, has been of continuous intrigue to 
both botanists and horticulturists. Although during the 
past 175 years several attempts have been made to clarify 
the various species that comprise this genus, no satis- 
factory compendium is presently at hand. It was Lind- 
ley who first produced a kind of monograph, or rather a 
revision of the genus in 1841, in his Sertum Orchidaceum 
under t. 48 with 101 species, and an expanded version 
of it in 1855 in his Folia Orchidacea with 198 species. 
These treatments were in use almost uninterruptedly 
until 1922 when anew attempt was made by Kraenzlin. 
It is unfortunate that the Kraenzlinian version was any- 
thing but an improvement on the former treatments. 
Since that time two synopses have appeared, one in 1970 
by Garay (Taxon 19: 443-467, 1970), and the other 
jointly by Garay and me in 1974 (Bradea 1(40): 893- 
424, 1974). IL believe that in this latter synopsis we have 
clearly outlined the infrageneric divisions which I intend 
to follow section by section in this series. These papers 
will not follow the sequence outlined in the key to the 
sections, but rather will treat them randomly depending 
on the demand and need of immediate attention. Conse- 


* Research Fellow, Orchid Herbarium of Oakes Ames. 


[ 183 ] 


quently this first study deals with the Section Cucullata. 

The actual identity of the plants with colorful and 
attractive flowers of the Section Cucullata have been 
plagued with confusion and misunderstanding since the 
early days of their introduction to European horticul- 
ture. During the nineteenth century hundreds of these 
plants were flowered in England and Europe and many 
of their floral variations were noted. For the most part, 
the early botanists considered the morphological differ- 
ences as being minor and not worthy of specific rank. 
Lindley, for example, chose to combine several different 
plants as varieties of O. cucudlatum including his earher 
described O. nubigenum. This action, since no univer- 
sally accepted Code of Nomenclature existed at that 
time, prompted Reichenbach to remark (Gard. Chron. 
367, 1867) “°... he (Lindley) called this plant a good 
species (nubigenum) but when he was older it had come 
into fashion to combine very heterogeneous types into 
one species because there were too many species’. De- 
spite Reichenbach’s attempt to separate these plants into 
distinct entities, the practice started by Lindley con- 
tinued and O. Phalaenopsis, O. oltvaceum as well as O. 
andigenum were declared to be varieties of O. cucullatum 
by Nichols in 1886, Veitch in 1892, and Gower in 1899, 
respectively. 

Reichenbach maintained a vigorous separation of the 
species in this affinity, attested by his drawings and de- 
scriptions. In 1922, Kraenzlin, in his monograph of On- 
cidium, attempted to expand the work of Reichenbach 
by establishing the Section Cucullata as a distinct unit. 
This effort, although noteworthy, included several spe- 
cies which belong to other affinities. He also omitted 
O. andigenum which he placed ina section together with 
QO. cornigerum tor reasons known only to him. In general 
Kraenzlin well understood his Section Cucullata, but he 


[ 184 | 


was rather careless and inconsistent in the allocation of 
the species. 

The circumscription of a section composed of seem- 
ingly similar flowering plants in itself is not a difficult or 
tedious task, but the delimitation of the species assigned 
to a given section is altogether a different matter. This 
latter approach demands a careful examination of all type 
specimens along with the associated descriptive data. 
Living material must be studied in the field and in cul- 
tivation, and compared with the types. Camera lucida 
drawings of the floral parts of type specimens as well as 
of additional material must be prepared for comparative 
purposes. Data presented in this paper, and those to be 
published in the future, are assembled through these 
methods. 

The plants of the Section Cucullata are endemic to the 
Andes of South America and grow in the cold and humid 
climate found at about 8,000 meters. Their range is from 
Venezuela southward to northern Peru, with an obvious 
concentration in Keuador. In the mid-nineteenth cen- 
tury, several collectors shipped plants by the thousands 
to England and Europe, which formed the basis for many 
of the herbarium specimens. These plants are relatively 
common in living collections today, although as, ex- 
pected, in rather small numbers, for repeated field trips 
to previously lucrative areas vividly tell of the encroach- 
ment by man. There is no doubt that these plants will 
continue to be found, but certainly not in the presently 
accessible areas known to current collectors. 

Because of the rather peculiar relationship between the 
‘allus of the lip and base of the column, I entertained 
the thought that the Section Cucullata should be ex- 
cluded from the genus Oncidium. In all of the species, 
the base of the column has a tumid member on either 
side projecting forward and enclosing the base of the 


[ 135 ] 


‘allus. Most of the species of the other sections of On- 
cidium have a short, variously tumid column-base, but 
only the Cucullata types enjoy the horseshoe-like shape 
which shelters part of the callus. Yet, analogous situa- 
tions exist in the Section Stellata, which is a different 
type of modification of the basic pattern. Until we have 
a satisfactory key to the various genera of the Oncidium 
alliance, I decided to maintain this section under 
Oncidium. 

In the past, several Odontoglossum species of the O. 
rigidum relationship have been referred to the Section 
Cucullata because of the connate lateral sepals and the 
sessile sepals and petals. A study of all the characters has 
necessitated the exclusion of these species from the genus 
Oneidium tor the above listed characters document an 
obvious case of parallel evolution rather than similarity 
based on close relationships. Likewise Oneidium Storhki 
and O. Warscewiczi must also be excluded from the 
Section Cucullata due to the different columnar struc- 
tures of the flowers. 

Several noteworthy characters within the section serve 
to distinguish the species. Of the 19 species recorded in 
this paper, ten have a tuft of hairs at the base of the 
elongate callus, while the remainder have a short tumid 
‘allus without hairs. With respect to the calli, three 
species—O. andigenum, O. mimeticum and O. sanguino- 
lentum—have five subglobose tubercles in three rows. 
All others are with three-membered, variously merged, 
tumid or keel-like structures. The familiar long nose-like 
tubercle is found in O. eucullatum, O. olivaceum, O. 
Kennedy and O. rhodostictum; long, sulcate callus which 
is tridenticulate at the apex, on the other hand, is limited 
to O. spathulatum, O. tripterygium and O. Dayanum. 

Five species—O. cucullatum, O. mimeticum, O. oliva- 
ceum, O. rhodostictum and O. sanguinolentum—have a 


prominently cucullate clinandrium, while the remainder 
have only a low collar which partially encircles the anther. 

The lip is the most conspicuous part of the flower in 
each species and contributes greatly to the overall size. 
Most species have a variously constricted lip with a well- 
defined isthmus or with a cuneate base to the midlobe 
suggesting a trend toward an isthmus. Otherwise, the 
midlobe is sessile and cordate, precluding an isthmus as 
in O. nubigenum. 

The most important non-floral character within the 
section is found in the inflorescence. Only O. cucullatum, 
O. Kennedyi and O. oltvaceum have a long, erect, rigid 
peduncle with a fractiflex rachis, while all others have a 
flaccid, arcuate peduncle with a sinuously flexuous rachis. 
This character alone is sufficient to separate the three 
species mentioned from the rest in the section. 


Section Cucullata A7vz/. in Pflanzenr. Heft 80: 
128: 1922. 

Type: Oneidium cucullatum Vind). 

The flowers range between 2 to 4 cm. vertically and 
have attractive color patterns. The sepals and petals are 
subsimilar, ovate-oblong, sessile, acute, and often con- 
cave. The color is variable, frequently mottled, often 
solidly colored with a well-defined margin. The lateral 
sepals are variously connate, always shorter than, and 
hidden by, the lip. The lip is conspicuous, usually con- 
stricted in the middle, with or without a well-defined 
isthmus; the side lobes subquadrate to rounded and often 
deflexed. The midlobe is spreading, emarginate, vari- 
ously maculated, occasionally on both sides. The crest 
consists of an odd number of keels or tumors with or 
without a hirsute base. The column is short, glabrous, 
occasionally with a cucullate clinandrium, and a thick- 
ened base; the rostellum is short. 


[ 137 ] 


The plants are rather small with aggregate or approxi- 
mate ovoid to pyriform and slightly compressed pseudo- 
bulbs which are enveloped by several imbricating sheaths 
of which the uppermost may be leaf-bearing. The one or 
more terminal leaves are linear to elliptical or narrowly 
lanceolate and subcoriaceous. The inflorescence, fre- 
quently precocious, emerges laterally from the basal 
bracts and is usually slender, arching, few-flowered, and 
racemose; occasionally it is rigid and erect, or many- 
Howered and paniculate, often conspicuously fractiflex. 


Key to Species 


Part of callus enclosed by the auriculate base 
of column adorned with a tuft of hairs . . . . . . . . . 2 


_ 


la. Part of callus enclosed by the auriculate base 


of column without a tuft of hairs . . . . . . . . 2. +. . 158 
8 Inflorescence rigid, erect, with more or less 
flexuous rachis . . . 0.0.0. ee ee ee ee SB 


Ya. Inflorescence flaccid, arcuate with sinuous 

PCN a ke RRS es ee RO eS ee 
3. Column with erect clinandrium, callus of lip 

keel-like, one-fourth to one-sixth of entire 

length of lip, triangular in lateral view, with 

a small, divaricately spreading lobe resembling 

nostrils on each side « « 6% 6 @ 6 se eee ee we & 
3a, Column with dorsally reclining clinandrium ; 

callus of lip keel-like, one-third of entire 

length of lip, triangular in lateral view consist- 

ing of three, parallel, distinct ridges of which 

the median is longer and nose-like without 

small spreading lobes on each side. . . . . 2. 2. 2 we OD 
4, Inflorescence racemose, rarely with a few short 

branches; sepals at least 16 mm. long; lip 30 

mm. long... 0... ee ee ee ee ee OF olivaceum 

var. olivaceum 

4a, Inflorescence paniculate with numerous, long, 

diffused branches; sepals up to 10 mm. long; 

lip up to20mm.long. . . . . . . 2 2. . . OW olivaceum 

var. giganteum 


[ 138 ] 


6a, 


8a. 


io} 


10a, 


Ll, 


lla. 


Column with distinct, linear processes on each 

side of the stigma; disc of lip with scattered 

wart-like papillae in center. . . Wee ew . OJ Kennedyi 

Column without any processes on each side of stigma; 

dise of lip without any wart-like papillae in center .... 6 

Flowers medium size; sepals not more than 

15 mm. long; lateral lobes of lip oblong- 

rectangular with obliquely truncate apex . . . O. cucullatum 
var. cucullatum 

Flowers large; sepals at least 20 mm. long; 

lateral lobes of lip never oblong-rectangular . 


Lateral lobes of lip rectangularly spreading 
obliquely triangular or dolabriform . . . . .  O. cucullatum 
var. dolabratum 
Lateral lobes of lip recurved toward midlobe, 
obliquely ovate. 2. 2... 2 ww ee ee OW cucullatum 
var. macrochilum 
Callus of lip elongate, consisting of three 
variously confluent, parallel ridges . 


&% 


Callus of lip short, consisting of five confluent 

tubercles in three rows, two on each side and 

the fifth in the middle. . ..........2.2.2. 2.18 
Petals unicolored except margins; clinandrium 

cucullate; callus of lip compressed, keel-like, 

the median ridge much longer than the very 

obscure lateral ridges, nose-like laterally . . . O. rhodostictum 
Petals marmorate to barred; clinandrium 

collar-like, marginate; callus of lip not com- 

pressed, the median ridge often considerably 

shorter than the lamella-like lateral ridges, 

never nose-like . . . .. . ae . ee eae ae 
Lateral sepals connate into a bifid sy aoa 

petals froma cuneate base elliptic-lanceolate, 

abruptly acute to acuminate .............421 
Lateral sepals free at least to middle, spread- 

ing; petals broadly elliptic, sessile, obtuse . . O. spathulatum 


Callus of lip obovate in outline, concave with 

entire, raised margins, bidentate at apex and 

with a shorter keel in the center originating 

from the middle of the callus . . . . . . . OJ Phalaenopsis 
Callus of lip quadrate to oblong in outline, 

conver, consisting of three parallel ridges of 

which the two outer ones are obscurely lobu- 


[ 139 } 


12 


a 


12a. 


12. 


13a. 


14, 


14a, 


7a. 


18a. 


19a. 


20. 


late, and the third in the center originating 
from the base of the lip between the column- 


WINGS: 2 6 oo Se 


Median keel much shorter than the beens ones 


Median keel as long as the lateral ones . 
Column slender; clinandrium low, 


clinandrium high, cucullate 


Column stout; 
Flowers rather small: lip when expanded 
distinctly pandurate, i.e., 
middle; midlobe of lip truncate to cuneate 
OCDASE 4 ce Se ee Ee 

Flowers medium to large, lip when expanded 
never pandurate, but constricted above the 


middle; midlobe of lip transversely reniform, 


constricted in 


deeply cordate at base . 

Lip when flattened truncate to subcordate at 
BOR % ee & 6 4 Re eS ee ee eS 
Lip when flattened cuneate at base 


Midlobe of lip sessile, transversely elliptic 
to cordate-reniform ew we ee 
Midlobe of lip cuneate to cuneate-flabellate 
Sepals and petals conspicuously muricate- 
papillose dorsally, especially towards base; 
petals distinctly mucronate at the more or 
less emarginate apex... ae ae 
Sepals and petals glabrous ee oe petals 
neither mucronate nor emarginate at apex 
Lateral lobes of lip ovate-oblong, forming an 
acute to subacuminate, open sinus with 
median lobe; callus large, basally produced 
into a retrorse, conical point . . . . . . . 
Lateral lobes of lip subrotund to obliquely 
obovate with rounded lobes, often imbricating 


the base of median lobe ; callus small without 


a retrorse, conical point . ........ 


Flowers medium size; lip 15-17 mm. long; 
callus quadrate in outline, porrect, composed 


of three ridges . . . . .. ee 


Flowers large: lip 22-24 mm. eee eallus 
transversely rectangular-oblong, distinctly 
curved upwards in front without any ridges 
but slightly depressed. . .....25.-. 


Petals with transverse bars; lateral lobes of 


(140 | 


marginate . 


12 

0. Dayanum 
O. tripterygium 
O. andigenum 
14 


O. sanguinolentum 


O. mimelicum 


O. azuayense 


O. nubigenum 


. O. aequinoctiale 


O. alticola 


O. tarquiense 


lip quadrate, truncate; midlobe of lip with a 

rectangularly transverse, erose apex . . . . . QO. erosilabium 
20a. Petals without transverse bars; lateral lobes 

of lip rounded; midlobe of lip 2-lobulate 


without an erose apex oo a oe ey -21 
21. Pseudobulbs one-leaved ; lip wider than long, 

transversely elliptic in outline . . . . . . O. chimborazoénse 
21a. Pseudobulbs two-leaved ; lip as long as wide, 

rhombic in outline . 2... . . . . . . . OF. tunguraguense 


SYSTEMATICS OF SPECIES 


Oncidium olivaceum H.B.K., Nov. Gen. et Sp. PI. 

1: 847, 1816. 

Syn. : Oneidium cucullatum subvar. olivaceum (H.B.K.) 
Veitch, Man. Orch. PI. pt. 8: 31, 1892. 

Oncidium cucullatum var. olivaceum (H.B.1K.) Gower 

in Garden 55: 88, 1899. 

Type: Colombia, Depto. de Cauca, Puracé near Pop- 
ayan. Coll. Humboldt no. 2020! (P). 

Observation: Dorsal sepal 16 mm. long, 6 mm. wide; lateral sepals 
cymbiform, bifid, 16 mm. long, 7 mm. wide; petals 15 mm. long, 8 
mm. wide; lip 3-lobed, lateral lobes obliquely ovate, 12 mm. long, 
7 mm. wide, midlobe from a narrow isthmus reniform, deeply cordate 
at base, bilobed in front, whole lip 30 mm. long, 32 mm. wide across 
midlobe. The callus is triangular laterally, keel-like, with a small, 
divaricately-spreading lobe, resembling nostrils on each side. 

Oncidium olivaceum is readily separable from O. cucullatum by its 
callus which is one-fourth to one-sixth of the length of the entire lip 
and by the presence of nostril-like lobes on each side of the nose-like 
keel, It is noteworthy that I have not seen any material of this spe- 
cies outside the Cauca area in Colombia. 


Oncidium olivaceum var. giganteum ( Hort.) Stacy, 

comb, nov. 

Basionym: Oncidium cucullatum var. giganteum Hort. 
in Garden 22: 166, 1882. 

Type: Reported only from cultivation; no specimen 
is known to exist. The original color plate is regarded 
here to represent the holotype. 


[ 141 ] 


Syn. : Oneidium olivaceum var. Lawrenceanum Gower 
in Garden 52: 26, 1897. 

Neotype: Ecuador, without exact locality. Coll. 
Lehmann no. 10081! (x, AMES). Since no type speci- 
men has ever been designated, the above-cited collection 
by Lehmann is selected here because duplicates of it al- 
ready exist in various herbaria under this name. 

Observation: The long, almost straight branches produced in pro- 
fusion lend a very distinct appearance to the plants of this variety. 
I have seen specimens from the Imbabura province of Kcuador and 
from the Cauca, near Popayan in Colombia. When more information 
will be available of this remarkable plant, it may prove to be a dis- 
tinct species. 

Illustrations : Cogniaux, Dict. leon. Orch. Oncidium pl. 28, 1899, as. 
O. cucullatum; Amer. Orch. Soc. Bull. 37: 13, 1968, as O. cucullatum. 


Oncidium Kennedyi Stacy, sp. nov. 

Type: Ecuador, along road from Quevedo to Lata- 
cunga, ca. 6000 ft. alt. Coll. G. Kennedy s.7./( AMES). 

KE. piphytica, caespitosa, usque ad 65 cm. alta; radici- 
bus crassiusculis, glabris: pseudobulbis ovoideis, bifoli- 
atis, usque ad 7 cm. altis; foliis lineari-oblongis, usque 
ad 22 em. longis, 2.5 cm. latis; inflorescentiis arcuatis, 
paniculatis, usque ad 65 cm. longis; pedunculo rigidius- 
culo, tereti; rhachide fractiflexa; sepalo postico valde 
concavo, anguste ovato-oblongo, acuto, 15 mm. longo, 
5 mm. lato; sepalis lateralibus inter se connatis, bifidis, 
ovato-cymbiformibus, 16 mm. longis, 7 mm. latis: pe- 
talis reflexis, ovato-ellipticis, undulatis, obtusis, 15 mm. 
longis, 6 mm. latis; labello 8-lobo, lobis lateralibus ob- 
lique ovatis, obtusis, lobo intermedio cuneato-flabellato, 
antice bilobulato, margine undulato, disco sparse papil- 
loso, callo lateraliter compresso, obscure 38-carinato; 
columna erecta, auriculis oblongo-linearibus Juxta stigma 
ornata; clinandrio suborbiculari, crenulato; ovario pedi- 
cellato 8.5 cm. longo. 


[ 142 | 


Prare NNVIII 


O. olivaceum 


O. olivaceum 


var. giganteum 


Observation: In many respects both the plants and flowers belong- 
ing to this new species resemble those of O. cucul/atum Lindl. It is 
unique in the whole alliance on account of the presence of distinct 
ears on each side of the stigma as well as the papillose lip. No doubt 
that many more undescribed species and forms are yet to be reported 


in this alliance. 


Oncidium cucullatum Lindl., Sert. Orch. sub. t. 21, 
1858. 
Type: Keuador: On the western declivity of Pichin- 
cha. Coll. : Jameson s.n./ (IX). 


Observation; The plants belonging to this species are rather small- 
flowered. Dorsal sepal 12 mm. long, 6 mm. wide; lateral sepals deeply 
concave, connate, bifid, 13 mm. long, 8 mm. wide; petals elliptic, 
obtuse, 12 mm. long, 8 mm. wide: lip 3-lobed, lateral lobes quad- 
rate, midlobe froma narrow isthmus reniform, deeply cordate, bilobed 
in front, 20 mm. long, 15 mm. across lateral lobes, 25 mm. across 
midlobe. The callus consists of three, keel-like, parallel ridges, the 
median ridge is triangular in lateral view, nose-like and longer than 
the lateral ridges which divaricate at front. 

The callus of the lip in the typical form as well as in the recognized 
varieties is always one-third of the length of the entire lip. Most 
probably because the type belonged to the Hooker Herbarium and 
because Lindley did not make a copy of it for himself, he confused 
it with O. sanguinolentum and with others now kept separate under 
O. olivaceum and O. mimeticum. 

Oncidium cucullatum is essentially Ecuadorian in distribution, but it 
may extend into Southern Colombia across the political boundary, up 


to Pasto in Depto. de Narino. 


Oncidium cucullatum var. dolabratum Stacy, 
var, NOV. 

Type: Ecuador, without proper locality. Coll. Jame- 
son s.n.! (IX). 

A var. cucullatum, floribus multo majoribus et lobis 
lateralibus labelli oblique triangulari-dolabriformibus 
differt. 

Dorsal sepal oblong-lanceolate, acute, 20 mm. long, 
6 mm. wide: lateral sepals connate, bifid, acute, 22 mm. 
long,7 mm. wide; petals ovate-lanceolate, acute, 19 mm. 


[ 144 ] 


PLATE X XIX 


. cucullatum 


var. dolabratum 


LAG 


[ 145 ] 


long, 10 mm. wide; lip 8-lobed, lateral lobes obliquely 
triangular-dolabriform, acute, 10 mm. long, midlobe 
from a narrow isthmus reniform, cordate at base, bilobed 
in front; callus an elevated, laterally compressed, nose- 
like keel, with a tuft of hairs at base, 10 mm. long; whole 
lip 30 mm. long, 22 mm. across lateral lobes, 80 mm. 


> 


across terminal lobe. 


Oncidium cucullatum var. macrochilum Lindl., 

Folia Orch. Oncidium 22, 1855. 

Syn.: Oneidium olivaceum var. macrochilum (Lind].) 
Sander, Orch. Guide 187, 1901. 

Type: Keuador, Quitinian Andes, on trunks of tree, 
at 4000 m., Coll. Jameson s.n./ (IXK-L, AMES-GH). 


Observation: Dorsal sepal narrowly elliptic, abruptly acute, 18 mm. 
long, 8 mm. wide; lateral sepals connate, bifid, deeply navicular, 20 
mm. long, 10 mm. wide; petals broadly ovate-elliptic, obtuse, 17 
mm. long, 12 mm, wide; lip 3-lobed; lateral lobes obliquely ovate, 
rounded, recurved toward midlobe; midlobe from a short isthmus 
reniform, cordate at base, bilobed in front; whole lip 28 mm. long, 
20 mm. across lateral lobes, 38 mm. across terminal lobe. 


Oncidium rhodostictum Krz]. in Pflanzenr. Heft. 
80: 2385, 1922. 
Type: Specimen without proper locality, cultivated 
by Sander s.n./ (W). 
Syn.: Oncidium cucullatum var. maculosum Laindl., 
Foha Orch. Oncidium 22, 1855. 
Oncidium olivaccum var. maculosum (Laindl.) Sander, 
Orch. Guide 188, 1901. 
Lectotype: Colombia, near Mariquita, in the forest 
of Quindiu. Coll. Linden 1887! (K—L), 7 hoe loco. 
Oncidium cucullatum var. purpurascens Hort. in Gar- 
den 21: 94, 1882. 
Type: Reported only from cultivation; no specimen 


is known to exist. 


[146 ] 


Prater XXX 


O. rhodostictum 


O. andigenum 


LAG 


[ 147 ] 


Oncidium cucullatum var. Chestertont Hort. in Garden 

23: 574, 1888. 

Type: Reported only from cultivation; no specimen 
is known to exist. 

Observation: Sepals 18 mm. long, 5 mm. wide; petals 18 mm. 


long, 7 mm. wide; lip 20 mm. long, 10 mm. across lateral lobes, 20 
mm. across terminal lobe. 

In appearance, this plant is very similar to O. mimeticum in growth 
habit as well as in the floral structure, especially the proportions of 
the lip. Yet, the two species can easily be separated because of the 
configuration of the callus. In O. rhodostictum, the callus of the lip is 
always nose-like, triangular in lateral view. In O. mimeticum, the callus 
is always a five-parted tubercle which is rectangular in lateral view, 


but never nose-like. 


Oncidium spathulatum (Lindl.) Stacy, comb. nov. 

Basionym: Oneidium cucullatum var. spathulatum 
Lindl., Folia Orch. Oncidium 22, 1855. 

Syn.: Oneidium nubigenum var. spathulatum (Lindl. ) 
Rehb.f. in Gard. Chron. 539, 1872. 

Oncidium cucullatum subvar. spathulatum (Lindl. ) 

Veitch, Man. Orch. Pl. pt. 8: 31, 1892. 

Type. Ecuador, without proper locality. Coll. Lobb 
s.n./ Veitch sale 1847 (IK—L). 

Oncidium Phalaenopsis var. excellens Linden in Lind- 

enia 12:58, t. 558, 1897. 

Lectotype: Ecuador, without precise locality. Im- 
ported and flowered by Linden s.n. No specimen is 
known to exist. The illustration in Lindenia is chosen 
here for the Lectotype. 


Observation: Dorsal sepal 16 mm. long, 7 mm. wide; lateral sepals 
connate to middle, 17 mm. long, 6.5 mm. wide; petals broadly el- 
liptic, obtuse, more or less mucronate, 14 mm. long, 8.5 mm. wide; 
lip constricted above middle with open, rounded sinuses, up to 25 
mm. long, 18 mm. across lateral lobes, 22-32 mm. across terminal 
lobe. 

Illustrations: Warner & Williams, Orch. Album 2; t. 96, 1883, as 
O. Phalaenopsis; Veitch, Man. Orch. PI. pt. 8: 30, 1892, as O. cucul- 
latum var. Phalaenopsis; Amer. Orch. Soc. Bull. 27: 758, 1958, as 


[ 148 ] 


PLateE XXXI 


O. spathulatum 


O. Dayanum 


[ 149 ] 


O. Phalaenopsis, Amer. Orch, Soc. Bull. 42: 1009, 1973, as O. 
Phalaenopsis. 


Oncidium Phalaenopsis Linden & Rchb.f. in Gard. 
Chron. 416, 1869. 
Syn. : Oncidium cucullatum var. Phalaenopsis (Linden 
& Rehb.f.) Nichols., Hl. Dict. Gard. 2: 485, 1886. 
Oncidium olivaceum var. Phalaenopsis (Linden & 
Rehb.f.) Sander, Orch. Guide 188, 1901, in syn. 
Type: Ecuador, without precise locality. Coll. Wallis 
sn! (W). 
Observation: Dorsal sepal 18 mm. long, 9 mm. wide; lateral sepals 


connate, bifid, 18 mm. long, 7 mm. wide; petals acute, 17 mm. long, 
10 mm. wide; lip 33 mm. long, 22 mm. across lateral lobes, 32 mm. 
across terminal lobe. 

The callus of the lip is very characteristic, obovate in outline, con- 
cave with entire raised margins, bidentate at apex, and with a third, 
shorter keel in the center originating from the middle of the callus. 
There is a good colored drawing from the type plant among Day’s 
sketches at Kew, Book 13, page 33, prepared in 1868, a year earlier 
than Reichenbach’s description. 


Oncidium Dayanum (Rchb.f.) Stacy, comb. nov. 

Basionym: Oncidium cucullatum var. Dayanum 
Rehb.f. in Gard. Chron. 834, 1871. 

Type: Ecuador, without proper locality. Imported 
and cultivated by Day, s.n./ (W). 

Syn.: Oncidium Phalaenopsis var. Brandtiae Hort. in 
Sem. Hortic. 2: 175, 1898. 

Type: Reported from cultivation, no specimens known 
to exist. The photograph published subsequently under 
O. Phalaenopsis in Orchid Rev. 9: 169, 1901, most proba- 
bly represents a picture of the type. 

Observation: Oncidium Dayanum is much smaller than O. triptery- 
gium. Dorsal sepal narrowly elliptic, acute, 13 mm. long, 6 mm. wide; 
lateral sepals connate, bifid, 16 mm. long, 7 mm. wide; petals sub- 
spathulate, elliptic, acute, 13 mm. long, 7 mm. wide; lip 3-lobed, 
wider than long, constricted above the middle, without an isthmus; 
whole lip 20 mm. long, 16 mm. across lateral lobes, 24 mm. across 


terminal lobe. 


| 150 | 


Oncidium Dayanum has been imported together with O. Phalaenopsis 
by Linden. Excellent color drawings exist at Kew among Day’s draw- 
ings, Book 15, page 56, drawn on Febr. 16 and 28, 1871. 

Illustrations: Amer. Orch. Soc. Bull. 27: 729, 1958; ibid. 42: 
1009, 1973; ibid. 44: 580, 1975, as O. Phalaenopsis; Dodson & Gilles- 
pie, The Biology of the Orchids, Frontispiece, 1967 as O. Phalaenopsis. 


Oncidium tripterygium Rchb.f. in Bot. Zeit. 10: 
694, 1852. 
Type: Ecuador, Loja. Coll. Warscewicz s.2./ (W). 
Observation: Dorsal sepal 18 mm. long; lateral sepals connate, 
bifid, 18 mm. long, 6 mm. wide; petals from a cuneate base elliptic- 
lanceolate, acute, 17 mm. long, 8 mm. wide; lip longer than wide, 
33 mm. long, 22 mm. across lateral lobes, 30 mm. across terminal lobe. 
The type material consists of a single flower in a rather poor state 
of preservation. In size it is very close to O. Phalaenopsis, but the 
callus of the lip is very different. It is quadrate to oblong in outline, 
convex, consisting of three parallel ridges of which the two outer ones 
are obscurely lobulate, the third in the center originates from the base 
of the lip between the column-wings and as long as the lateral ones. 
Illustration: Il]. Hortic. 17: t. 8, 1870, as O. Phalaenopsis. 


Oncidium andigenum Linden & Rchb.f. in Gard. 

Chron. 416, 1869; ibid. 539, 1872. 

Syn.: Oneidium cucullatum var. andigenum (Linden 
& Rehb.f.) Hort. in Garden 22: 166, 1882. 

Oneidium cucullatum subvar. andigenum (Linden & 

Rchb.f.) Veitch, Man. Orch. PI. pt. 8:31, 1892. 

Oneidium  olivaceum var. andigenum (Linden & 

Rehb.f.) Sander, Orch. Guide 184, 1901. 

Type: Ecuador, without proper locality. Coll. Wallis 
s.n./ (W). 

Observation: Dorsal sepal 6 mm. long, 2.5 mm. wide; lateral se- 
pals connate, bifid, cymbiform, 7 mm. long, 3 mm. wide; petals elliptic- 
oblong, obtuse, 6mm. long, 2.5 mm. wide; lip pandurate, 7 mm. long, 
4.5 mm. across lateral lobes, 5 mm. across terminal lobe. 

Although references have been made to this species in literature, 
I have seen only one collection in addition to the type to which I can 
satisfactorily refer here. There is a color drawing with details of flow- 


ers of this species in the Kew Herbarium among Day’s drawings, 
Book 18: page 54, drawn on October 19, 1874. The corresponding 


[151 ] 


specimen was sent to Reichenbach by Day, and it is mounted on the 
type sheet. Oncidium andigenum is the smallest flowered in the group 
and has a five-parted callus. 


Oncidium sanguinolentum (Lindl. ) Schltr. in Fedde 

Rep. Beih. 6: 98, 1919. 

Basionym: Leochilus sanguinolentus Wind. in Bot. 
Reg. 80: Misc. p. 91, 1844. 

Syn.: Oneidium cucullatum var. sanguinolentum 
(Lindl.) Lindl., Folia Orch. Oncidium 22, 1855. 

Oncidium olivaceum var. sanguinolentum (Lindl. ) San- 

der, Orch. Guide 188, 1901. 

Type: Colombia, La Guayra. Coll. Barkers.n./ (KX-L). 

Observation: The type specimen consists of two separately mounted 
flowers and a colored drawing apparently prepared froma living flower 
by Lindley. The flowers are rather small: dorsal sepal 11 mm. long, 
5 mm. wide; lateral sepals 13 mm. long, 5.5 mm. wide; petals 11 
mm. long, 6 mm. wide; lip 15 mm. long, 10 mm. across base, 14 
mm. across the bilobed terminal part. 

The reduction of this species by Lindley to O. cucul/atum is difficult 
to understand because of the distinctions in growth habit and the 
differently constructed callus. Oncidium sanguinolentum flowers from an 
undeveloped new growth and has a 5-parted callus: while O. cucul- 
latum flowers from mature pseudobulbs and has a nose-like, keeled 
callus. 

Illustrations: Dunsterville and Garay, Venezuelan Orchids Illustr. 
5: 220, 1972, only Fig. B, as O. andigenum. Foldats in Lasser, Flora 
Venez. Orch. 15 (5): 360, 1970, as O. nubigenum: Bertha Ospina, 
Orquideas de la Clarita, 75, 1972, as O. cucullatum. 


Oncidium mimeticum Stacy, sp. nov. 

Type: Columbia, Depto. Norte de Santander, Ocana. 
Coll. Schlim no. 411! (IKK-L). 

Epiphytica, caespitosa, variabili; radicibus flexuosis, 
glabris; pseudobulbis approximatis, ovoideis vel pyri- 
formibus, vulgo unifoliatis, cataphyllis obtectis; foliis 
anguste oblongo-ellipticis vel lanceolato-oblongis, obtu- 
sis, basi conduplicatis, quam inflorescentiis vulgo brevior- 
ibus; inflorescentiis elongatis, prominenter pedunculatis, 


Led 


PLATE XA XATI 


O, sanguinolentum 


LAG 


supra laxe paucifloris: floribus vulgo secundis; bracteis 
late triangularibus, acutis, pedicellis multo brevioribus ; 
sepalo postico elliptico, obtusiusculo, valde concavo, 
usque ad 15 mm. longo, 6 mm. lato; sepalis lateralibus 
inter se fere usque ad apicem in synsepalo connatis, 
cochleato-cymbiformibus, apice divaricatim bidentatis, 
usque ad 16 mm. longis, 7 mm. latis; petalis late ellip- 
ticis vel obovatis, obtusiusculis, reflexis, usque ad 13 mm. 
longis, 8 mm. latis; labello distincte lobulato, lobis later- 
alibus auriculatis vel subquadratis, lobo intermedio trans- 
verse reniformi, bilobo, antice vulgo sinuato vel promi- 
nenter exciso, basi valde cordato, margine undulato, disco 
tuberculis 5-nis, cariniformibus elevatis ornatis: toto la- 
bello usque ad 25 mm. longo, 30 mm. lato, basin inter 
lobos laterales 12 mm. lato; columna humili, 6 mm. 
alta; clinandrio cucullato; ovario pedicellato ca. 15-20 
mm. longo. 


Observation: The rather abundant material of this species present 
in the various herbaria has been confused repeatedly with O. cuculla- 
tum Lindl. The specimens belonging to this new species can readily 
be differentiated by the shape of the callus which consists of 5 tuber- 
cles or humps arranged in three rows; 2 humps on each side and a 
fifth one in the middle which isshorter than the lateral ones. Although 
the structure of the calli is similar to O. sanguinolentum ( Lindl.) Schltr., 
the flowers of the latter species differ, in addition to being much 
smaller, in the construction of the lip which is pandurate and the 
midlobe of lip is truncate to cuneate at the base. 

The original error dates back to Lindley who confused this appar- 
ently distinct species with O. cucu//atum in Paxton’s Flower Garden. 
It should be noted that the plate in Paxton’s Flower Garden is based 
on an abnormal specimen, showing a terminal inflorescence. 

Illustrations: Paxton Fl. Gard. 3: t. 87, 1852, as O. cucullatum; 
Fl. des Serres 8: t. 853, 1853, as O. cucullatum; ibid. 23: t. 2457, 
1880, as O. cucullatum; Veiteh, Man. Orch. Pl., pt. 8: 30, 1892, as 
O. cucullatum; Amer. Orch. Soc. Bull. 27: 750, 1958; ibid. 44: 581, 
1975, as O. cucullatum; Dunsterville & Garay, Venez. Orch. Ill. 5: 
220, 1972, Fig. A only, as O. andigenum; Foldats in Lasser, Flora 
Venez. Orch. 15(5): 360, 1970, dotted outline of lip only, as O. 


nubigenum., 


[ 154 ] 


XXXITI 


PLATE 


g 
= 
s 
ra 
£ 
E 
S 


Oncidium mimeticum var. flavidum (B.S. Wms.) 

Stacy, comb. nov. 

Basionym: Oncidium cucullatum var. flavidum B.S. 
Whis., Orch. Grow. Man. ed. 4, 229, 1871. 

Syn.: Oncidium cucullatum var. flavidum Linden, Cat- 
alogue under nos. 88 to 86, 1870. 

Oneidium olivaceum var. flavidum (B.S.Wmms. ) Sander. 

Orch. Guide 187, 1901. 

Neotype: Since no material is known to exist of the 
original introduction by Linden in 1870, Catalogue nos. 
33 and 36, the watercolors prepared by Day of the flow- 
ers sold under no. 38 and 86, and which are now kept at 
Ikew are regarded here as representing the holotype. 
Although this unusual color variant was noticed at first 
by Linden, the plant became officially described only by 
Williams in 1871. The color variant is maintained here 
distinct on account of its flowering time which is in De- 
cember. The flowering period of O. mimeticum is April— 
May. 


Illustration: Ill. Hortic. 25: t. 305, 1878, as Oncidium cucullatum. 


Oncidium azuayense Krzl. in Pflanzenr. Heft 80: 
3821, 1922. 
Type: Ecuador, southern slope of the Azuay moun- 
tains. Coll. Lehmann no. 587! (G). 


Observation: Dorsal sepal broadly elliptic, obtuse to truncate, oc- 
casionally somewhat retuse at mucronate apex, dorsally sparsely but 
distinetly papillose-muriculate, 9 mm. long, 5 mm. wide; lateral sepals 
connate, bilobulate in front, the inside margins of lobes often imbri- 
cate, dorsally prominently keeled and sparsely papillose-muriculate, 
especially near base, 10 mm. long, 8 mm. wide; petals broadly ellip- 
tic, rounded to deeply emarginate at the mucronate apex, dorsally 
with a few papillae near the base, 10 mm. long, 7 mm. wide; lip 
3-lobed, lateral lobes oblong to subquadrate, midlobe sessile, cordate- 
reniform, bilobed in front, 16 mm. long, 14 mm. across lateral lobes, 
24 mm. across terminal lobe; callus with a retrorse median tooth. 

The plants of this rather commonly occurring species in the Azuay 
region can easily be recognized by the dorsally papillose-muriculate 


[ 156 | 


PLATE XXXIV 


O. nubigenum 
LAG 


sepals and petals and the frequently deeply emarginate petals with 
a mucronate tip. The sepals and petals are always fleshy-coriaceous. 

Illustrations: Amer. Orch. Soc. Bull. 27: 748, 1958, as O. nubi- 
genum; ibid. 27: 757, 1958; Florida Orchidist 10: 14, 1967, as O. 


nubigenum. 


Oncidium nubigenum Lindl., Gen. and Sp. Orch. 
Pl. 197, 1833. 
Syn.: Oneidium cucullatum var. nubigenum (Lindl. ) 
Lindl. Folia Orch. Oncidium 22, 1855. 
Oncidium olivaccum var. nubigenum (Lindl.) Sander, 
Orch. Guide 188, 1901. 
Type: Ecuador, on the ridge of Azuay mountains. 
Coll. Jameson .s.2/ (K,K-L). 


Observation: Dorsal sepal elliptic, dorsally carinate, 11 mm. long, 
6 mm. wide; lateral sepals, connate basally to } of their length, cari- 
nate toward apex, 13 mm. long, 5 mm. wide; petals elliptic, obtuse 
to abruptly subapiculate, never mucronate, 1] mm. long, 7 mm. wide; 
lip 3-lobed, lateral lobes semiovate, without a sinus and imbricating 
the reniform midlobe, 17 mm. long, 13 mm. across lateral lobes, 23 
mm. across terminal lobe; callus very large in comparison with others 
of the immediate relationship, with a retrorse conical point. 

The flowers of O. nubigenum can be compared only with those of 
O. azuayense, but the two differ from one another in many important 
points. The flowers in O. nubigenum are of thin texture, sepals and 
petals never papillose dorsally, petals never mucronate, lateral sepals 
free almost to base, and the callus of the lip is robust. The flowers in 
O. azuayense are rather coriaceous in texture, sepals and petals papil- 
lose dorsally, petals mucronate and commonly emarginate, lateral 
sepals connate nearly to apex, and the callus of lip is insignificant. 

What prompted Lindley to reduce this species to a variety of O. 
cucullatum in 1855 will never be understood. But, it surely has confused 
the issue ever since. Already in 1867 Reichenbach complained (Gard. 
Chron. p. 376) * . when the lamented Dr. Lindley was a young 
lynx-eyed observer, he called this plant a good species. When he was 
older, and it had come into fashion to combine very heterogeneous 
types in ‘one species” because ‘there were too many species’, he be- 
lieved it was his duty to cancel some of his own, and this Oncidium 
was degraded to the rank of a variety. We believe this was a mistake, 
since the many flowers... . never show any cucullate anther-bed, 
never have a nasiform keel on the lip, nor is there ever an isthmus 
to the lip’. 


[ 158 | 


Notwithstanding Reichenbach’s clear analysis of the distinctions, 
Lindley’s error has been carried on to the present. Veitch in his 
Manual of Orchidaceous Plants pt. 8: 30, 1887, suggests that since 
O. nubigenum was described first, Lindley should have reversed the 
reduction by calling O. cucullatum a variety of O. nubigenum. This 
opinion is echoed by Dodson and Frymire in Amer. Orch. Soc. Bull. 
27: 750-751, 1958, by calling O. cucullatum and O. Phalaenopsis a 
subspecies of O. nubigenum. Fortunately, no legitimate transfers were 
made at that time. 


Oncidium alticola Stacy, sp. nov. 

Type: Ecuador, Prov. Azuay, Nudo de Potate, pass 
between headwaters Rio Tarqui and Gir6n, ca. 9,000 ft. 
alt. ‘‘Kpiphyte. 5 segments pale to ochraceous-yellow 
with purplish tinge; 6th white with purple splotch at 
base. Base of column purple, apex white. ‘Bulbs’ (pseu- 
dobulbs) called ‘guagra saccha’ and when ground said to 
be diuretic; but natives often eat them when thirsty.” 
Coll. Camp no. EK-2163! (AMES). 

E;piphytica, caespitosa, usque ad 16 em. alta; radici- 
bus flexuosis, glabris; rhizomate abbreviato, crasso ; pseu- 
dobulbis approximatis, lateraliter compressis, anguste 
cylindraceis vel cylindraceo-ovoideis, 2-foliatis, usque ad 
4 cm. altis: foliis lineari-oblongis vel oblongo-oblanceo- 
latis, acutis, usque ad 11 cm. longis, 1.2 cm. latis; in- 
florescentiis, satis gracilibus, remote paucifloris, usque 
ad 16cm. longis; bracteis ovato-cucullatis, acutis, 4mm. 
longis; sepalo postico obovato-oblanceolato, acuto, 12 
mm. longo, 4 mm. lato; sepalis lateralibus inter se usque 
ad medium connatis, anguste obovatis, acutis, 12 mm. 
longis, 4 mm. latis; petalis ellipticis, obtusis, breviter 
apiculatis, 10 mm. longis, 6 mm. latis: labello 4-lobo, 
lobis lateralibus subrotundis, inter se 9 mm. latis, lobo 
intermedio e cuneata basi flabellato, antice valde bilobo, 
lobis subrotundis, disco ima basi callo 8-partito, tumido 
ornato, toto labello 19 mm. longo, antice 17 mm. lato; 
columna humili; clinandrio tenuiter marginato; ovario 
pedicellato usque ad 2 cm. long. 


[ 159 ] 


Observation: The large lateral lobes of the lip and the cuneate- 
flabellate midlobe readily identify the plants belonging to this new 
species. It appears to be rather common in the Azuay region. 

The following additional specimens belong here; Ecuador: Azuay- 
Tarqui. Coll. Harling & Andersson no. 18205 bis! (AMES, S); Cuena, 
Coll. Strobel s.n./ (AMES); without exact locality. Coll. Rodrigo 
Escobar-R. no. 1380! (AMES). 

Illustrations: Amer. Orch. Soc. Bull. 32: 905, 1963, as O. nubi- 


genum., 


Oncidium tarquiense Stacy, sp. nov. 

Type: Ecuador, Province Azuay, Portete del Tarqui, 
Cuenca-Girén, 2700-2950 m. altitude. Coll. Harling & 
Andersson no. 18205! (S). 

Epiphytica erecta, usque ad 30 cm. alta; radicibus 
flexuosis, glabris, pseudobulbis ascendentibus ovoideis 
vel pyriformibus, 2-foliatis, basi vaginis chartaceis ob- 
tectis, usque ad 5 cm. longis, 1.5 cm. latis: foliis satis 
tenuibus, lineari-oblanceolatis, obtusis, usque ad 10 cm. 
longis, 1.5 cm. latis: inflorescentiis gracilibus, paululo 
arcuatis, laxe secundifloris: racemo laxifloro: bracteis 
infundibuliformibus, oblique ovato-triangularibus, 5mm. 
longis; sepalo postico elliptico, obtuso vel subacuto, 16 
mm. longo, 6 mm. lato; sepalis lateralibus quarta parte 
basali inter se connatis, supra liberis, anguste ellipticis, 
acutis, 17 mm. longis, 5 mm. latis; petalis late ellipti- 
cis, acutiusculis, 14 mm. longis, 8 mm. latis; labello 
3-lobo, lobis lateralibus oblique ovatis, lobo intermedio 
maximo, cuneato-flabellato, antice bilobulato, lobulis 
rotundatis; disco basin callo minuto, subquadrato, sur- 
sum curvato ornato; toto labello 24 mm. longo, 28 mm. 
lato; columna humili; clinandrio marginato; ovario 
pedicellato usque ad 85 mm. long. 

Observation: The flowers in general appearance are larger than 
those found in O. a/ticola. The transversely rectangular-oblong callus 


which is distinetly curved upwards is unique in the whole alliance, 
hence affords an excellent field character for identification. 


[ 160 } 


PLtatreE XXXV 


O. alticola 


O. tarquiense 


LAG 


Oncidium aequinoctiale Stacy, sp. nov. 

Type: Ecuador, without precise locality. Coll. André 
s.n.f (K). 

I. piphytica, usque ad 80cm. alta; radicibus flexuosis, 
glabris; pseudobulbis approximatis, anguste ovoideis, 
unifoliatis, usque ad 5 cm. altis; foliis lineari-lanceolatis, 
acutis vel subacuminatis, usque ad 17 cm. longis, 1.5 
cm. latis; inflorescentils erectis, supra racemosis, laxe 
paucifloris, usque ad 80 cm. longis; floribus satis tenui- 
bus, secundis; bracteis cucullatis, acutis, ovariis pedicel- 
latis multoties brevioribus, 4mm. longis; sepalo postico 
valde concavo, elliptico, obtuso, 11 mm. longo, 6 mm. 
lato: sepalis lateralibus ad tertiam partem apicalem inter 
se connatis, valde concavis, obtusis, usque ad 12 mm. 
longis, 6 mm. latis; petalis obovatis, obtusis, inconspicue 
apiculatis, 11 mm. longis, 7 mm. latis; labello 3-lobo, 
lobis lateralibus divergentibus, ovato-oblongis, lobo in- 
termedio sessili, paululo cuneato, transverse elliptico, 
antice bilobo, utrinque sinuoso; callo satis prominenti, 
3-partito, partitione intermedia retrorse conica; toto la- 
bello 18 mm. longo, basin 14 mm. lato, antice 24 mm. 
lato: columna humili, clinandrio marginato. 


Observation: In floral structures the plants of this new species re- 
semble those of O. nubigenum. It differs from O, nubigenum especially in 
having free lateral lobes forming open, acute to acuminate sinuses with 
the midlobe of the lip; the midlobe is cuneate not cordate at the base. 


Onsidium erosilabium Stacy, sp. nov. 

Type: Ecuador, without precise locality. Cultivated 
by Rodrigo Escobar-R. no. 1879! (AMES). 

K.;piphytica erecta, usque ad 30 cm. alta; radicibus 
flexuosis, glabris; pseudobulbis ovoideis, lateraliter com- 
pressis, usque ad 6 ecm. altis; foliis linearibus, acutis, 
18-25 cm. longis; inflorescentiis gracilibus, paululo arcu- 
atis, paucifloris, usque ad 10 cm. longis; floribus albidis, 
sepalis petalisque pallide brunneo maculatis striatisque, 


[ 162 | 


PiatE XXXVI 


O. aequinoctiale 


QO. erosilabium 


LAG 


[ 163 ] 


labello basin purpureo maculato, callo aurantiaco; sepalo 
postico elliptico, obtuso, 11 mm. longo, 6 mm. lato; 
sepalis lateralibus tertia parte basilari inter se connatis, 
oblongo-ellipticis, acutis, 14 mm. longis, 5 mm. latis;: 
petalis late ellipticis, obtusis, 11 mm. longis, 6 mm. 
latis: labello e cuneata basi 8-lobo, lobis lateralibus sub- 
quadratis, truncatis, lobo intermedio sessill, transverse 
reniformi, antice rectangulariter late eroso: disco venis 
incrassatis ornato; callo cylindrico, antice 8-partito; toto 
labello 18 mm. longo, 21 mm. lato; columna erecta; 
clinandrio marginato; ovario pedicellato usque ad 24 
min. longo. 

Observation: In floral structure the plants of this species are simi- 
lar to O. chimborazoénse, but the barred petals and the squarely erose 
lip differentiates it immediately not only from that species but from 
every member of the whole alliance. I have seen another specimen 
referable to this species in the Kew Herbarium; it was cultivated 
during the turn of the century under the name of O. cucul/atum var. 


Oncidium chimborazoénse Stacy, sp. nov. 

Type: Ecuador, Chimborazo, 3200-8400 m. altitude. 
Coll. F.C. Lehmann no. 215! (AMES, Kk). 

K.;piphytica, ascendenti, usque ad 25 cm. alta; radici- 
bus flexuosis glabris ; pseudobulbis approximatis, oblique 
ovoideis, unifoliatis, usque ad 25 mm. altis: foliis per- 
gameneis, anguste lanceolato-ellipticis, subacutis vel ob- 
tusis, usque ad 18 cm. longis, 1 cm. latis; inflorescentiis 
erectis vel paululo arcuatis, supra laxe paucifloris; flori- 
bus ut videtur secundis; bracteis infundibuliformibus, 
ovato-triangularibus, acuminatis, 5 mm. longis; sepalo 
postico obovato-elliptico vel elliptico-oblanceolato, acu- 
to, vel abrupte subacuminato, usque ad 12 mm. longo, 6 
mim. lato; sepalis lateralibus basin inter se breviter con- 
natis, anguste ellipticis, acutis vel subacuminatis, usque 
ad 18 mm. longis, 5 mm. latis; petalis late ellipticis, 
obtusis, usque ad 11 mm. longis, 6 mm. latis; labello e 
cuneata basi 8-lobo, lobis lateralibus obliquis, apice ro- 


[ 164 | 


PLATE AXXVIIT 


O. chimborazoénse’ 


O. tunguraguense 


LAG 


tundatis; lobo intermedio maximo, transverse reniform], 
antice valde bilobulato, basi subecordato, disco basin callo 
tumido, 3-partito ornato, toto labello usque ad 23 mm. 
longo, 28 mm, lato; columna humili, crassa, clinandrio 
marginato: ovario pedicellato usque ad 25 mm. longo. 


Observation: Oncidium chimborazoénse is larger than O. tunguragu- 
ense, especially in the size of the flowers. It differs from the latter in 
the plants having only one-leaved pseudobulbs and the lip is much 
wider than long. 

There is another specimen at Kew collected also in Chimborazo by 
Cross no. 52! (K), who sent it to the Veitch establishment in 1882. 

Illustration: Bot. Mag. 94: t. 5708, 1868, as O. cucullatum var. 
nubigenum; Belg. Hortic. 19: t. 19, 1869, as O. nubigenum; Dict. Icon. 
Orch. Oneid. t. 30, 1900, as O. nubigenum; Amer. Orch. Soc. Bull. 
41: 1001, 1972, as O. nubigenum. 


Oncidium tunguraguense Stacy, sp. mov. 

Type: Keuador, Vulcan Tunguragua, 8000-8400 m. 
alt. Aug.-Sept. Coll. F.C. Lehmann no. 8572! (Ix). 

Epiphytica, erecta, usque ad 25 cm. alta: radicibus 
flexuosis, glabris; pseudobulbis approximatis, pyriform- 
ibus, lateraliter compressis, bifoliatis, usque ad 4 cm. 
altis; foliis ellipticis vel lanceolato-ellipticis, acutis, usque 
ad 13 cm. longis, 2.5 cm. latis; inflorescentils erectis, 
sursum arcuatis, laxifloris, usque ad 25 cm. longis; flori- 
bus ut videtur secundis; sepalo postico oblanceolato vel 
obovato-oblanceolato, acuto vel obtusiusculo, usque ad 
10 mm. long, 4.5 mm. lato; sepalis lateralibus supra 
basin inter se breviter connatis, lineari-oblongis, acutis 
vel obtusiusculis, 11 mm. longis, 8 mm. latis; petalis 
ellipticis obtusis vel abrupte acutis, usque ad 9 mm. 
longis, 5 mm. latis: labello in ambitu trapezoideo, 3- 
lobo, lobis lateralibus e cuneata basi triangulis, lobo inter- 
medio transverse reniformi, antice retuso vel subbilobulo : 
disco venis incrassatis ornato; callo tumido, 8-partito: 
toto labello usque ad 15 mm. longo, 16mm. lato; colum- 
na humili; clinandrio marginato; ovario pedicellato usque 


ad 2 cm. longo. 
| 166 | 


Observation: From the closely related O. chimborazoénse this new 
species is readily distinguishable by the 2-leaved pseudobulbs and the 
rhombic lip which is approximately as long as wide. The sepals and 
petals are maroon inside, much paler and suffused outside; the lip is 
white with heavy crimson blotches on the basal half especially on the 
lateral lobes; the callus is orange. 


ACKNOWLEDGMENTS 


The steady cooperation of my colleagues at the Orchid 
Herbarium of Oakes Ames, and especially the guidance 
of Dr. Leslie A. Garay, have been a continuous inspira- 
tion throughout the preparation of thisstudy. Their help 
is deeply appreciated. My special thanks to Dr. Peter 
Taylor, the Orchid Herbarium, Royal Botanic Gardens 
and to Dr. Luciano Bernardi, Conservatoire et Jardin 
Botaniques, Geneve, for their kind cooperation in pro- 
viding specimens and drawings of types. For permission 
to reproduce their color slides so essential to this study 
I am indebted to Mr. Gilberto Escobar-R, Mr. Rodrigo 
Escobar-R, Dr. George Kennedy and Mrs. Henry T. 
Northen. The publication of color plates was made possi- 
ble through the generosity of Mr. Maurice Tl’. Freeman, 
Mr. Walter Hunnewell, Mr. and Mrs. Lee B. Kuhn, 
the Lear Siegler Foundation, Mrs. Henry 'T. Northen, 
Mr. Robert M. Scully, Sr., Mr. Richard Seifert and 
Mr. Robert G. Stone. Their thoughtfulness and help 
are sincerely acknowledged. ‘To Miss Gertrude Ahern 
I am grateful for the dedicated attention given by her 
to this paper. 


[ 167 ] 


PLATES 


[169 | 


Top left: Oncidium cucullatum. Photograph: R. Northen. 

Top right: Oncidium cucullatum. Photograph: G. Escobar-R. 
Bottom left: Oncidium olivaceum. Photograph: G. Escobar-R. 
Bottom right: Oncidium rhodostictum. Photograph: G. Kennedy. 


[170] 


PLATE XXXVIIL. 


Top left and right: Oncidium Kennedyi. Photographs: G. Kennedy. 
Bottom: Floral details of Oncidium Kennedyi. 


XXXIX. 


PLATE 


Top: Oncidium mimeticum. Photograph: R. Northen. 
Bottom: Oncidium mimeticum. Photograph: G. Kennedy. 


[174] 


PLATE XL. 


Top left: Oncidium mimeticum. Photograph: G. Kennedy. 

Top right: Oncidium sanguinolentum. Photograph: G. Escobar-R. 
Bottom left: Oncidium sanguinolentum, Photograph: R. Escobar-R. 
Bottom right: Oncidium sanguinolentum. Photograph: G. Escobar-R. 


[176 | 


PLATE XLI. 


Top left and right: Oncidium mimeticum. Photographs: G. Kennedy. 
Bottom: Oncidium tripterygium. Photograph: R. Northen. 


[178] 


PLATE XLII. 


[179] 


Top: Oncidium Phalaenopsis. Photograph: R. Northen. 
Bottom: Oncidium Dayanum. Photograph: R. Northen. 


[180] 


XLII. 


PLATE 


[181] 


Top left and right: Oncidium Dayanum. Photographs: G. Kennedy. 
Bottom left and right: Oncidium spathulatum. Photographs: G. Kennedy. 


[182] 


PLATE XLIV. 


[183] 


Top: Oncidium tarquiense. Photograph: J. Stacy. 
Bottom: Oncidium alticola. Photograph: R. Escobar-R. 


[ 184} 


PLATE XLV. 


[185 | 


Top and bottom: Oncidium nubigenum. All photographs by G. Kennedy. 


[186 | 


PLATE XLVI. 


[187] 


Top left: Oncidium alticola. Photograph: G. Escobar-R. 
Top right: Oncidium erosilabium, Photograph: R. Escobar-R. 
Bottom: Oncidium chimborazoénse. Photograph: G. Kennedy. 


[188] 


PLATE XLVIL. 


[189] 


Top: Oncidium aequinoctiale. Photograph: R. Northen. 
Bottom: Oncidium tunguraguense. Photograph: J. Stacy. 


[190] 


PLATE XLVIII. 


[191] 


APR 19 197 


CAMBRIDGE, MassAacHusETTs, APRIL 9, 1976 VoL. 24, No. 8 


PLANTAE COLOMBIANAE XIX 
E PARTIBUS AMAZONICIS WITOTORUM PLANTAE 
FRUCTUARIAE SATIVAE NOVAE 
BY 
RicHarp Evans SCHULTES 


In studiis ethnobotanicis meis de plantis alimentis in- 
diorum witotorum e Amazonia colombianae duas arbores 
novas inveni: unam e genere leguminosarum, Inga; al- 
teram e genere apocynacearum, Macoubea. In regionibus 
Rio Karaparand et Rio Igaraparana utraeque arbores ab 
indigenis coluntur. 

Doctori Thomas S. Elias pro descriptione speciei In- 
gae novae gratias ago. Similiterque erga Patrem Miguel 
Junyent, Matrem Holga Edith Sajona et Sororem Ligia 
Maria Ochoa, propagatores doctrinae christianae catholi- 
cae in colonia San Rafael apud Rio Karaparana inferio- 
rem, pro beneficentiis suis, praecipue mihi flores ex 
arbore typica praebuisse, gratus sum. 


APOCYNACEAE 


Macoubea witotorum PR. FE. Schultes spec. nov. 

Arbore usque ad quadraginta ped. alta, dense foliosa, 
ex radice singulare truncis multis, in terranon inundabile 
culta; cortice laeve, crasso, in maculis albido-griseo, 
abundanter albo-lactifere. Folia valde chartacea, supra 
atroviridia, infra pallide viridia, late elliptica, base ae- 
qualia, apice obtusa vel rarenter subacuta, lamina 20-24 


[ 193 ] 
ECONOMIC BOTANY LIBRARY 


OF OAKES AMES 
HARVARD BOTANICAL MUSEUM 


PuatTE XLIX 


(fi Yi, 


LA 


\Ss 


MACOUBEA t) 
witotorum #.€. Schu/tes ki JBC 


1, leafy branch, approximately } natural size. 2, inflorescence, ap- 
proximately 5 natural size. 3, flower, approximately 14 natural size. 
t, basal portion of flower dissected, approximately twice natural size. 
5, stamen, approximately 4 times natural size. 6, corolla lobe, ap- 
proximately 4/5 natural size. Drawn by Josuua B, Crank 


PuatE L 


MACOUBEA 
witotorum 
ae R.E. Schultes 


B, dissected fruit, showing 


A, fruit, approximately s natural size. 
9 ’ pp . ri 
placement of seeds, approximately 4 natural size. C, seed, approxi- 


mately natural size, with detail of surface greatly enlarged. 
Drawn by Josuvua B. Cirark 


em. longa, 11-15 em. lata, utrinque glabra, nervis utrinque 
prominentibus; petioli 1-2.5 cm. longi. Inflorescentiae 
corymbosae, multiramosae, usque ad trigintiflorae, bracte- 
atae, minutissime griseo-puberulae, axibus principalibus 
robustis, usque ad 8 cm. longis, pedicellis brevibus, pler- 
umque I cm. vel brevioribus, minute bracteolatis. Flores 
albido flavi vel eburnei, non fragranti, plus minusve 25 
mm. longi. Calyx persistens; corollae lobis vix contor- 
tis, base glandulosis, ovatis, 15 mm. longis et basin 10 
mm. latis: corollae tubo quam lobis multo breviori, in 
fauce albo-piloso; antheris sessilibus, sagittatis, usque ad 
4 mm. longis; ovario dense albo-piloso, stylo 0.8-1.2 
mm. longo annulo crasso atrobrunneo coronato et crista 
quinquelobata capitata in acumine 0.8 mm. producto. 
Apocarpium magnum, usque ad 16 cm. longum, 11 ¢m. 
<9 cm. in diametro, irregulariter reniforme vel stomach- 
iforme, petioli basin versus submammiforme, extus in 
maturitate brunneum vel subaureo-brunneum, sordide 
granuloso-squammulosum, intus viridulum praeter sem- 
inarum lectum atrobrunneum, usque ad 15 mm. crassum, 
pulpa brunneola liquida dulce ; seminibus plurimis, rubro- 
brunneis, curvatim elliptico-oblongis, superficie dense et 
grosse vermiculato-foveolatis, inter foveas minutissime 
eristatis, cristis magnopere minute indumento § albo- 
spongioso indutis, 16 mm. longis, 6 mm. latis; pedunculo 
lignoso, 10 mm. crasso, usque ad 13 cm. longo, calyce 
valde persistente. 

Cotompra: Comisaria del Amazonas, Rio Karaparana, San Rafael 
(in vicinity of El Encanto), near confluence with Rio Putumayo. “Tree 
up to 40 ft. Many trunks from root. Cultivated. Bark smooth, light 
grey with whitish patches. Latex copious, white, sticky. Leaves 
thick, coriaceous. Outside of fruit brown; inside light green except 
where seeds lie (which is brown). Pulp liquid, light brown, sweet, 
eaten. Seeds dark brown. Seeds and pulp from one fruit 250 ce. 
Witoto name: 00-rroo’-soo-je (in Rio Karaparana), o0-koo-je (in Rio 
Igaraparana)’’. March 80-April 2, 1970. Richard Evans Schultes 26072 
(Tyveus in Herb. Gray: Typr Dupticati in Keon. Herb. Oakes Ames ; 


| 196 | 


PiLaTtTeE LI 


Fruit of tree from which the type material of Macoubea witotorum 
was taken. Photograph: R.E. ScHuites 


[ 197 | 


Herb. Nac. Colomb.).—Same locality. ‘““Flowers yellow-cream or 
whitish yellow, without fragrance. Taken from type tree’’. August 
1970. P. Miguel Junyent sine num. (Fioris Typrin Herb. Gray ; Econ. 
Herb. Oakes Ames: Herb. Nac. Colomb. ). 


99 
. 


Genus Macoubea, ut hodie comprehenditur, duas spe- 
cies includit: MM. guianensis Aubl., quae totam terram 
regionem guianarum atque Brasiliae amazonicae inhabi- 
tat, sed varietas eius, MW. guianensis var. pubiflora Mona- 
chino, non nisi ex loco typi prope civitatem Iquitos, e 
regione Amazoniae peruvianae observatur: Macoubea 
Spruce’ (Muell.-Arg.) Markgraf, cuius typus a Richard 
Spruce apud Rio Uaupés juxta Brasiliae Colombiaeque 
terminum collectus est, tantum in regione Rio Negro 
Brasiliae amazonicae afferitur; distributio varietatis eius, 
Macoubea Sprucei var. paucifiora (Spr. ex Muell.-Arg. ) 
Monachino, cum specie congruit. 

Secundum Monachino (Lloydia 8 (1945) 296), ‘‘spe- 
cies Macoubearum inter sese arcte affines. Differentiae 
primae quae in elementis variis discernendae possint in 
florium proportionibus magnitudinibusque fundantur, 
sed nequaquam constantes sunt’. 

Macoubea witotorum a speciebus adhoc cognotis non 
solum florium characteres proportionesque differt, sed 
etiam a JM. guianense fructus formae proportionesque 
valde dissimilis est. 

Praeterea haec species nova ab utraque specie florium 
characteribus minoribus distingui potest. Macoubea wito- 
torum corollae lobi 15 mm. longi atque 10 mm. lati 
metiuntur, dum in MM. guianense 3-4 mm. et in M. 
mim. lati sunt; corollae tubus quam lobus 


Sprucet 5-7 
multo brevior qui in JZ. guianense quam lobus dimidium 
vel longior, et in JZ. Sprucet prope eadem aequilongus 
est. Macoubea witotorum a M. guianense fructus forma 
proportioneque facile distinguitur: in priore mericarpio 
immaniter reniforme vel stomachiforme est, 16 ¢m. longo 


[ 198 | 


Piarye E11 


a) 
Macoubea C PUY ANNENELE 


Illustration of Macoubea guianensis, published in Aublet: Plantes 
de la Guiane Francaise 2 (1775) t: S78. 


[ 199 ] 


et 1] cm. X 9 cm. in diametro metitur; in posteriore, 
globoso, circiter 6-8 cm. in diametro metitur. 

Secundum exiguos libros, Macoubea arbores sunt sil- 
vicola et semper sua sponte crescunt et silvas in terra 
humida sed supra inundationem annuam_ preoptans. 
Macoubea witotorum est arbor quae ut videtur solum in 
cultura exsistit et semper in terra alta, bene siccata, nun- 
quam inundata prosperat. 

Indigenae tribus witotorum ut arbores solum in regione 
Rios Karaparana et I garaparana coleri affirmant,et verum 
est ame nunquam ipsis in multis meis annis explorationis 
in regionibus amazonicis invenitis per quos annos cogita- 
tiones meas plerumque versus plantas cultas dirigi. Si 
naturaliter occurrit, ea est rara. 

Indigenae fructus parte terminale acutiore praecidunt 
vel in eo foramen faciunt et copiosam pulpam fulvam 
ducunt quae in forma liquidi dulcis adest. Infeliciter, 
studia nutrimenta huius liquidi non facta sunt, sed sine 
dubio sacchari abundantia saturata est. Semina non 
devoruntur. 

Arbores mensibus martio aprileque fructus abundanter 
ferunt. Unusquisque exempla quae in cultura vidi mul- 
tas (usque ad quinque vel sex) truncos corpulentos ex 
radice unico surgentes habuerunt. Utrum hoe proprium 
incrementum mutilationibus trunci primari an ab indi- 
genis eventus est non potuit, sed indigenae ipsae arborem 
normaliter hoe modo crescere affirmant et se nullo modo 
arborem ad conditionem faciendam truncare vel secare. 
Certe, hae incrementi consuetudo mensuram = arbore 
coronae Magnopere auget atque, propter hoc, abundan- 
tem segetem fructuum. 

Doctor C. Sastre (Museum d°‘Histoire Naturelle, 
Paris) specimina florescentia quae ut videtur esse elus- 
dem Macoubeae species in pago La Chorrera prope Rio 


Igaraparana colligit. 


[ 200 ] 


Pyare LE! 


DIAGONAL SHADING 
ON INSET INDICATES 
AREA SHOWN 
IN DETAIL BELOW 


lauitos f A 
R 1) LETICIA ! 5 


: XN 
“Si = 4 ~ rd au 
a Ao (CIARG : 


- 
Ya RETIRG ENTRE Rios Pa ieee 


& 


Re 


i ; c 
5 ( ws) 
(@) tt aie 
\ - y 
KARAPARANA-IGARAPARANA =, L 6 ~~ 
REGION US \ | Om iin 
of the ( ; 
COLOMBIAN AMAZON © . 


)  t 
AN fo * We AI) af if VA 
wa (c 4 p ae OO 


CotomBra: Comisaria del Amazonas, Rio Igaraparana, La Chorrera. 
**Arbre, 10 m., latex blane abondant, petales jaunes, fruits comest- 
ibles, pulpe liquide avee graines noires. Nom witoto: ukurai.’’ Sep- 
tember 29, 1973. C. Sastre 2340. 


LEGUMINOSAE 


Inga chorrerana Elias spec. nov. 

Arbores parvae; ramis teretibus, sparse ferrugineo- 
tomentellis, lenticellis conspicuis. Ramuli ferrugineo- 
tomentosi. Folia petiolis 4-5 cm. longis, dense ferrugineo- 
tomentosis, partim inter Juga superiora foliolorum dila- 
tatis; ala late ellipticis vel obovatis, 6-12 mm. latis. 
Foliola 6-jugata, petiolulis 2-8 mm. longis, dense fer- 
rugineo-tomentosis; lamina elliptica, apice apiculata, 
basi rotundata, 5.5-15 em. longa, 2.8—5.5 cm. lata, sub- 
coriacea, nervis lateralibus 16-20, subtus conspicuis; 
glandulae rhachidis 38-8.5 mm. diam., orbiculares; stip- 
ules non visae. Inflorescentiae multiflorae, spiciformes, 
3-5 cm. longae, terminales aut in axillis superioribus 
dispositae; pedunculis 1-2 cm. longis, dense ferrugineo- 
tomentosis; bracteis late ovalis, apice acuminatis vel sub- 
apiculatis, 5-6 mm. longis, dense ferrugineo-tomentosis ; 
calyx cylindricus, sessilis, 7-9 mm. longus, sparse tomen- 
tosus, lobis ovatis 1-1.5 mm. longis; corolla cylindrica, 
apice vix dilatata, 1.7-1.9 em. longa, dense villosa, lobis 
ovatis 2.5-8 mm. longis; stamina tubo vix exserto; fila- 
menta alba, ad 8.5 mm. longa: antherae dorsifixae ; 
pistillum sessile, teres, ca. 2 mm. longum, glabrum. 
Legumen coriaceum, in maturitate areis ciliorum diffu- 
sis ferrugineo-villosum, plano-compressum, ad 45 cm. 
longum, 5-7 cm. latum, ca. 1 cm. crassum, marginibus 
vix elevatis. Semina sine testa, cotyledonibus 8-8.2 cm. 
longis, 1 em. latis. 

CoLtombra: Comisaria del Amazonas, Rio Igaraparana, La Chorrera. 
Small tree. Pulp of fruits edible. Cultivated. Fruit 1-14 ft., long, 


slender, thickness of index finger. Flowers white, fragrant. June 6, 
1942. Richard Evans Schultes 3896. (Tyrus in Herb. Gray). 


( 202 | 


PLatE LIV 


Habit drawing, approximately | natural size. Pod, approximately 
s natural size. Inflorescence, approximately natural size. 


Drawn by Irene Brapvy 


[ 203 ] 


Hance speciem arborem fructiferem ob siliquaram mag- 
narum pulpa albam dulcem semina cingens coleri ferunt. 

Doctor Elias seripsit: ‘“‘Jnga chorrerana in Sectione 
Inga, Series Spectabiles propter proprietates florales 
fructuosas debet. Ab aliis speciebus affinibus folio sex- 
jugati, folium apice apiculato atque fructu pubescente 
distingueri potest. Ad Ingam spectabilem Willd. propin- 
qua atque fortissime proxima ad J. Go/dmannu Pittier 
ut videtur est.” 


[ 204 ] 


PACHYTENE CHROMOSOME MORPHOLOGY 
AND ITS BEARING ON INTERSPECIFIC 
AND INTERGENERIC RELATIONSHIPS 

OF COIX* 
BY 
J. VENKATESWARLU', R.S.K. CoaGantr, 
AND PanuGant! N. Rao’ 


The genus Coix is a member of the tribe Maydeae 
(family Gramineae) which comprises seven genera. ‘Two 
of these, Zea and Tripsacum, are native to the New 
World, while the rest, Coix, Chionachne, Trilobachne, 
Sclerachne and Polytoca, are native to southeast Asia 
(Mangelsdorf and Reeves, 1939). The Maydeae is a taxo- 
nomic rather than a natural assemblage of genera (Cha- 
ganti, 1965), with monoecism as the feature in common to 
both the New World and Asiatic members. However, 
from time to time, the existence of possible phylogenetic 
relationships between the two geographically separated 
groups has been postulated (Anderson, 1945; W.C. 
Galinat, personal communication). 

Coix is the largest of the five maydeaceous genera na- 
tive to Asia. Its systematics is confusing (Venkateswarlu 


* Investigation supported by a grant from the Indian Council of 
Agricultural Research, New Delhi. 


‘Department of Botany, Andhra University, Waltair, Andhra 
Pradesh, India. 


*The New York Blood Center, 310East 67th Street, New York, 
N.Y. 10021. 


[ 205 | 


and Chaganti, 1973). At least three highly polymorphic 
species assemblages are recognized, namely C. aquatica 
Roxb., €. Lachryma-Jobi L., and C. gigantea Iwoen. 
C. aquatica and C. Lachryma-Joli have chromosome 
numbers of 2n=10 and 20 respectively, while in the case 
of C. gigantea populations with chromosome numbers 
of 2n=20 and 40 have been reported (Venkateswarlu 
and Chaganti, 1978). 

Comparative chromosome morphology, especially at 
the pachytene stage of meiosis, where the chromosomes 
are more extended than at mitotic metaphase and often 
present characteristic features, is a useful parameter in 
understanding interspecific and intergeneric  relation- 
ships. In the case of Coix, such studies have been few 
(Venkateswarlu and Chaganti, 1973). We have studied 
and report here the pachytene chromosome morphology 
in populations of Coix representing the three species. 


Materials and Methods 


Samples of seeds from populations representing the 
three species were assembled from different regions of 
India. Some were collected in the field by the authors, 
while others were collected for them by colleagues in 
different parts of the country. In addition, two samples 
from Japan and one trom Brazil (countries in which Coix 
is not native, but into which it has been introduced for 
use as a fodder, cereal or ornamental) were also obtained. 
The geographic origin and method of collection of all the 
seed material is listed in Table |. Plants were raised from 
these seeds in the experimental gardens of the Depart- 
ment of Botany of the Andhra University. Young in- 
florescences were fixed in 8:1 ethanol-acetic acid. After 
24-48 hours of fixation, the fixative was replaced by 70% 
ethanol. The material was stored in this fluid at 4° C, 
until used for study. 


r 


[ 206 | 


Acetocarmine squash preparations were made of young 
anthers. Appropriate nuclei were photographed, and 
camera lucida drawings were made of them from tempo- 
rary preparations. Chromosomes were measured from 
camera lucida drawings alone, and their lengths, which 
include the centromeric regions, are expressed in micron 
units. Numbers are assigned to chromosomes on the 
basis of decreasing order of length: thus chromosome 1 
is the longest of the complement. Individual chromo- 
somes are identified on the basis of their length, arm- 
ratio (length of long arm/length of short arm), and the 
pattern of distribution of heteropyenotic and eupycnotic 
region. 

Observations 


Coixv aquatica Roxb. The chromosomes of this species 
are difficult to study at pachytene, because they are long 
and often intertwined. In addition, the chromosomes 
possess long heteropyenotic segments which often ex- 
hibit non-homologous association (Plate LV), thereby 
making it difficult to follow the chromosome from one 
end to the other. Deep staining heteropycnotic regions 
are present in the proximal regions of both arms. Addi- 
tional heteropyenotic regions are present at other loca- 
tions on the chromosome arms. ‘The chromosome arms 
terminate in pronounced and dark staining chromomeres. 
Exact measurements of the heteropycnotic regions were 
not possible to obtain, because of variability in their ex- 
tent (see Venkateswarlu and Chaganti, 1973). Camera 
lucida drawings of the five chromosomes are represented 
in Plate LVI, while the idiogram represented in Plate 
ILX is based on mean values of length and arm ratio of 
five measurements for each chromosome (see Table II). 
The third longest chromosome of the complement has 
the nucleolus organizer in the long arm in a sub-terminal 
position (Plate LX). 


‘uMOUYUN 3d1NOS AleUul 

“lig ‘suapiey d1ue}0g A}ISI9AIU Ls) 

dy} UL Sain}[Nd wWoIs Saadeosa se 

pajiejs aaey pinoy ‘sndulesy AjisIaa 

-1UQ) BIYPUY 9} UO PIM BSulMmois 

_ sioyjyne 9y} Aq paqdaTjoo ~—s sjuryd Jo Auojoo ¥ ‘pyiMy sndwes 


Sl-o a89[[0D [einqpnousy ‘jedioutsg :Aq patjddns BIpU] ‘d10]eBquIIOZ 


(s]eyr) UIo}sey) eIpUy 
Je[nsuluag Ula}seayynos jo asuel 


3-5 (OMSY) stoyine vyj Jo suo Aq pa}da][OD UTeJUNOU! [B}SeOD ay} Ul LuIsejJUeUY 7 
LD umou yun uMmouyuUN 4 
Bipuy jo 
Gp-- AVAING [BOIURIOg ‘jsIuRJOg [eUOIZoy :Aq parddns BIpuy ‘Wessy ry 
eIpuy jo ‘WYRM VIIA] “eA 
BP--Q AvAInS [eoIuUe}Og ‘JsIuRJOg [eUuOIsoy :Aq patiddns eIpu] ‘Wessy 1QO[-DUAALYIDT “D 
8 WO (JMSY) Sssoyine ayi Jo auo Aq pa}daq[oo BIpu] ‘ysepeid e1ypuy “ 
uonerodi07 
te ¥)D Spaas jeuONeN “WweeYyd “A°D “Aq :Aq parddns BIpu] ‘essliC “ 
g-Y'D AllsiaalugQ 1esnes ‘apueg “YS ‘Jorg :Aq patddns BIpU] ‘ySepetg eAyupRly pojonbv “>D 
Jequiny uonisinboy jo apoyy UISLIQ sIydeIs0a5) xx S3100dS 
uoyeindog 


Io UOISSIDDY 


, UONVSNSIAUI SIY} UI pasn ‘UZ0Y VazuDTIG ‘Dy pue ‘"] 1gof-DuAAYIDT “DQ “GxOY vIIDNbHY “D “ZILA ‘X10d JO 


saidads 391Y} 9Y4] 0} Sulsuojeq suonjejndod snorea ay} JO [el1a}eW peas Jo uO}ISInboe Jo spoul pue UIsIIO D1ydeuso0ar) 


I ATaAVL 


[ 208 | 


‘untieqiaH 


AJISIOAIUL) RIYPUY JY} Ul paytsodap uveq aARY 9A0gP pajsl] SatjalieA puke satdads ay} [Je Jo SUsUIOAdS UINTIeqIOY x» 
‘sojdures paas yim sn parddns oy ¢ uuIN]OO UI pajsl] ajdoad snorieA ay} 0} [NJa}e13 Iv OM y 


eIpuy jo 

I°9D = Aaarng [eotuejog ‘ory *S eI]OY “Iq :Aq partddns 
99D (QYNd) szeyyne ay} Jo auo Aq pajoaT]oo 
ojnedg ors 

TD ‘orlewy ‘1S ‘lasneyyeeyos “AY “Aq :Aq paddns 
erpuy jo 


Ga)  Avains |eo1uej}Og ‘ory “Ss BI[OY “Iq :Aq parjddns 


oJOAY ‘AVISIOAIUL) OAYIES 

‘ainj[nosy jo Jusuijiedaq ‘A10}e10qe’] Bul 

€-d -peeig pue souaINs dolla ‘epeiey iq :Aq parjddns 
Bipuy jo 

2-2 AVAING [edIURIOg ‘jsIURJOg [eUOISay :Aq patjddns 
ojoAY ‘AJISIOAIUS) OAYIES 

‘gin}[Nosy jo Juawjiedaq ‘Ai1ojeloqe’y] Bul 

Il-D -paeig pue aouatog doi ‘eperey ‘iq :Aq patjddns 
eipuy jo 

qOI-D AeAINS [edIuRIJOg ‘JsTUR}Og [BUOIsay :Aq patjddns 


erpuy jo 
ROT-D AVAING [edIURIOg ‘JsIuRlOg [eUOIsay :Aq patddns 


(s}eys) U191S9M ) 
eIpu] JO Iejnsuluag U13}sOM 


-yjnos jo asuel ulejUNOW j[e}seo7 DIJUDSIS ‘dD 


“p91g] 

pue 4afij1Mow “SIA 

U99M}9q J}eIPIULI9} 

-UL AT[BOIBOTOYCI0J 

eIpU] ‘iseUeUYy 1g0[-DUUKAYIDT “ZD 


[Izelq « 


jdeis (Jaueuloy) uand-pu “IeA 
eIpu] ‘ulessy 1g0[-DUKAYIDT “9D 


uedef 


jdeys vdups0uajs “1eA 


eIpu] ‘Wessy 190[-DUKAYIDT “Dd 


uedef 


eIpuy ‘und “e1yaq “ 
YEM 4afipiuou “IBA 


eIpu ‘ung elyeq 190[-DUAAYIDT “D 


{ 209 ] 


Cow Lachryma-Jobi lL. Chromosomes of all the popu- 
lations examined, with the exception of one (C-8 in Table 
I), show differential staining into eu- and heteropycnotic 
regions as in the case of C. aquatica. Heteropycnotic 
segments are always present in the proximal regions of 
both arms. In this species also the chromosome arms 
terminate in deep staining chromomeres. In addition, 
terminal as well as intercalary knobs are present. The 
sizes and numbers of knobs vary in the different popu- 
lations. Thus, some populations are devoid of Knobs 
(e.g., population C-4b), while others possess them (e.g., 
eight knobs are present in population C-4a). Detailed 
studies were not performed of knob frequencies in the 
various populations. A complete analysis of the pachy- 
tene chromosome morphology was achieved in one popu- 
lation (C-8), in which the chromosomes do not exhibit 
differential staining (Plate LVII, a, b). An idiogram 
(Plate I,X) was constructed on the basis of average value 
of up to five measurements of lengths and arm ratios for 
sach chromosome (see ‘Table II). The sixth longest 
chromosome of the complement has the nucleolus organ- 
izer inthe long arm in a nearly terminal position (Plate 
ILX). Plate LVIIL illustrates the nucleolus organizing 
chromosome in a population (campus wild) that shows 
differential staining of chromosome regions. 

C. gigantea Koen. Of this species, populations have 
been reported by several investigators (Darlington and 
Janaki Ammal, 1945; Nirodi, 1955; Venkateswarlu and 
Chaganti, 1973) with chromosome numbers of 2n=20 
and 2n=40. The population that we studied (C.G. 1, 
which is native to the mountain range on the west coast 
of peninsular India called the Western Ghats) exhibited 
variation in chromosome numbers ranging from 2n=18 
to 2n=22. One of the plants with 18 chromosomes 
(which at meiosis forms nine bivalents) exhibited well 


tL 21: 


PLatE LV 


Portion of pachytene nucleus of C. aquatica showing non-homologous 


association of centromeres and heteropycnotic regions. 


[ 211 ] 


spread pachytene chromosomes (Plate LIX, a, b). All 
the chromosomes of this species, Just as in the case of 
those of C. aquatica and C. Lachryma-Joln, show differ- 
ential staining into eu- and heteropycnotic regions, the 
latter being present in the proximal as well as interstitial 
regions of chromosome arms. Furthermore, the chromo- 
some arms terminate in deep staining chromomeres. An 
idiogram constructed on the basis of mean values of at 
least ten measurements of lengths and calculation of arm 
ratios for each chromosome is presented in Plate LN 
(see Table I1). The eighth longest chromosome of the 
complement has the nucleolus organizer situated almost 
terminally on the entirely heteropyecnotic short arm 


(Plate LN). 
Discussion 


Variation in chromosome morphology and knob numbers 
in C. Lachryma-Jobi: The species of Coix are predomi- 
nantly outbreeding. Four to six varieties have been 
recognized in C. Lachryma-Jobi based on morphological 
features of the fruit case (Mimeur, 1951; Bor, 1960). 
The varieties interbreed readily if grown together and 
allowed to open pollinate and their identity is lost within 
a few generations. Hence, the varietal differences are 
based on gene mutations, and the varieties arose through 
geographic isolation of populations or through selection 
by man. The variation observed at the population level 
in the chromosome phenotype: namely, the degree of 
heteropyenosis in chromosome arms and the sizes and 
numbers of knobs; can also be considered part of the 
same evolutionary processes that lead to varietal deline- 
ation. A detailed study of the variation of the chromo- 
some phenotype has not yet been accomplished, but 
available information points to existence of a considerable 
amount of it. The fact that, in one of the populations 


[ 212] 


PLatTe LVI 


1\Op 


Chromosome 2 


Chromosome 1 


Chromosomes 4(A) and 5(B) Chromosome 3. “a, - 


Camera lucida drawings of the five chromosomes of C. aquatica at 
pachytene. 


| 2ts: | 


(C-8), the chromosomes did not exhibit heteropycnosis 
might indicate that this feature is in some way controlled 
by the genotype. It is well known that several facets 
of chromosome form and behavior are under genotypic 
control (Rees, 1961; Chaganti, 1965). Knobs occur on 
the pachytene chromosomes of other members of May- 
deae, e.g., Zea and Tripsacum. In these genera, num- 
bers and positions at which knobs occur are characteristic 
features of populations. In C. Lachryma-Jobi var. typica 
collected at Coimbatore, a city in southwestern India, 
Nirodi (1955) observed five terminal and one interclary 
knobs. Ina population (C-8) of the same variety collected 
at a different location (Anatagiri in the coastal mountain 
range in southern peninsular India called the Eastern 
Ghats), we found only two terminal knobs. In yet an- 
other population (C-4a) of the same variety originating 
from a different locality in India (Assam, a hilly province 
in eastern India), we found eight knobs. The relation- 
ship of knob variation to geographic distribution of the 
populations remains to be studied. 

Non-homologous associations of centromeres and hetero- 
pycnotic regions in C. aquatica: In C, aquatica, at pachy- 
tene, non-homologous centromeres as well as non- 
homologous heteropyenotic regions are frequently asso- 
ciated (Plate LV). These associations, however, fall apart 
before diakinesis. The chromosomes of C. Lachryma- 
Jobi and C. gigantea exhibited none of these characteris- 
tics. The significance of such association is not known; 
however, it is likely that exchanges might take place in 
the associated regions and lead to reciprocal transloca- 
tions as suggested by Venkateswarlu (1958). The genome 
of C. aquatica is characterized by a degree of instability ; 
complex translocations are of frequent occurrence in 
natural populations(Venkateswarlu and Chaganti, 19738). 

Comparison of the genomes of the three species: A com- 


[ 214 | 


> LVII 


‘E 


Puat 


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-Ayord ye (] [qe], ul g 


+s | -—_—_—H1 


aN 


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Q uorjeindod) vordh} IVs 1g0f-vuhiyov'y “2 JO 2aSd010dsosJo1W & Jo snaponu aq J, (®) 


( we $6 
7 4 


parison of the idiograms of the three species (Plate LX) 
reveals that, except for the nucleolus organizing chromo- 
some, the chromosomes of the three species are quite 
dissimilar in their morphology. The length of each 
chromosome, its arm ratio, and the contribution its 
length made to the total length of the genome are given 
for the three species in Table II. C. aquatica has the 
longest chromosomes of the complement, with a genome 
which is 401.9% long (mean chromosome length of 
80.38). The genomes of C. Lachryma-Jobi and C. gig- 
antea respectively are 582.0% (mean chromosome length 
of 58.24) and 488.84 (mean chromosome length of 
54.31) long; thus C. Lachryma-Jobi and C. gigantea 
have genomes of approximately equal length. They are 
also shorter than that of C. aquatica by about 40%. The 
nucleolus organizing chromosomes of the three species, 
even though of widely differing lengths and arm-ratios, 
are similar: they are median to submedian, and the nu- 
cleolus organizer occupies a sub-terminal position. The 
dissimilar morphology of chromosomes between the pre- 
sumably basic diploid C. aquatica and the derived poly- 
ploid species C. Lachryma-Jobi and C. gigantea must 
reflect, on the one hand, the effects of hybridity and 
chromosome doubling which presumably have been in- 
volved in the emergence of the species with higher chro- 
mosome numbers, and, on the other hand, the proneness 
of the genome of C. aquatica to re-patterning due to 
translocation (Venkateswarlu and Chaganti, 1973). Inter- 
specific hybrids between C. aquatica and C. Lachryma- 
Joli, C. Lachryma-Jobiand C. gigantea, and C. aquatica 
and C. gigantea are feasible, and the chromosomes in the 
I’, hybrids exhibit intergenomic pairing (unpublished ob- 
servations of Rao). Thus, at least one genome is common 
to all three species even though its presence cannot be de- 
tected by comparison of chromosome morphology alone. 


[ 216 ] 


Piate LVIII 


B 


(a) Portion of a pachytene nucleus of C. Lachryma-Jobi var. typica 
(population Campus wild) showing the nucleus organizer chromosome. 
This population exhibits differentinl staining of eu- and heteropcynotic 
regions of the chromosomes which is evident in this cell. (b) A camera 
lucida drawing of the nucleolus organizing chromosome. 


Comparison of the genomes of Cow, Sorghum and maize: 
The suggestion made by Anderson (1945) that maize 
originated in southeast Asia as an amphidiploid of a hy- 
brid between species of Coix and Sorghum possessing a 
diploid chromosome number of 10, though rejected on 
a number of grounds (Mangelsdorf and Oliver, 1951; 
Chaganti, 1965), makes interesting a comparison of the 
morphology of the chromosomes of C. aquatica and spe- 
cies of Sorghum with a diploid chromosome number of 
10 and those of maize. Pachytene chromosome mor- 
phology has been reported by other investigators in two 
species of Sorghum with a diploid chromosome number 
of 10: viz., S. intrans (Garber, 1947) and S. purpureo- 
sericeum (Reddi, 1958). The chromosomes of JS. intrans 
are uniformly dark staining, and the centromeres are 
hard to locate in them. Three pairs are of one length; 
the remaining two are shorter and participate in nucleolus 
organization. The chromosomes of S. purpureo-sericeum 
ure in the same size range as those of maize and possess 
dark staining chromomeres in the proximal regions of 
both arms, making the centromeres easy to locate. 
Neither species possess knobs. The genome of C. aqua- 
tica, at pachytene, is about 1.7 times longer than that of 
maize (data from this study compared to measurements 
of maize chromosomes at pachytene given in the publi- 
‘ation of Rhoades, 1955), and the chromosomes them- 
selves are quite dissimilar in morphology to those of 
maize. Chromosomes of C. aquatica show differentially 
stained eu- and heteropyecnotic regions. ‘They are also 
devoid of knobs. Maize chromosomes do not exhibit 
similar differential staining, and they possess knobs. 
Morphological comparisons, thus, do not reveal common 
features between the genomes of C. aquatica, S. intrans, 
and S. purpureo-sericeum on the one hand and maize on 
the other. This observation by itself, in the absence of 


[ 218 ] 


PLatE LIX 


<<] 


2s 
St 


e 


5s 
15 
e 
8s @ eae 


- 
a 
os 

- 


4s 


Pad 
-—" 


6 
4% 


15 
~ 


7s 
eee i 
7 
$i 
3s bad am 


ae | 


B 
(a) The nucleus of a microsporocyte of C. gigantea at pachytene. 
Differential staining of eu- and heteropycnotic regions is clearly seen. 


(b) A camera lucida drawing of the same nucleus. 5=short arm; 


L=long arm; open circle =centromere; broken circle = nucleolus. 


genome analysis based on hybridization, cannot be con- 
sidered evidence against possible existence of true phylo- 
genetic relationships between these species. As discussed 
earlier, even in the case of the genus Coix, comparison 
of the chromosome morphology of the three species does 
not indicate the presence of the common genome demon- 
strated by intergenomic pairing of chromosomes in in- 
terspecific hybrids. 

Chromosomes of C. Lachryma-Jobi and maize exhibit 
a few similarities. They fall in approximately the same 
length range at pachytene (data from this study com- 
pared to measurements of maize chromosomes given in 
the publication by Rhoades, 1955) as well as at diakine- 
sis and metaphase I (Longley, 1941). Knobs, the sizes 
of which and positions on chromosomes vary in different 
populations, are present in both species. There is, how- 
ever, no evidence based on genome analysis to indicate 
that these similarities reflect true phylogenetic relation- 
ships. The many attempts made at producing viable 
hybrids between maize and Coix have so far met with 
failure (Mangelsdorf and Reeves, 1989: Venkateswarlu, 
1963). 


Summary 


Lengths, arm-ratios, and other morphological features 
of chromosomes at the pachytene stage of meiosis are 
reported for three species of Coix, namely, C. aquatica 
(2n=10), C. Lachryma-Jobi (2n=20), and C. gigantea 
(2n=18). The genome of C. aquatica is the longest of 
the three species and is about 40% longer than that of 
either of the other two species. When stained with 
acetocarmine the chromosomes of C. aquatica show con- 
spicuous differential staining into eu- and heteropyenotic 
regions. They also exhibit non-homologous association of 
centromeres and heteropyenotic regions. C. Lachryma- 


[ 220 ] 


is 


PLATE 


‘IT 2148], Ul paquasaid soijzei-Wiie pue SYySUI] Jo San[eA UBITI Jo SISeq 94} UO 
paiedaid vajunsis “cy pure ‘igof-vuhsyovy *Q ‘voipnby *Zy Jo samosomoiyo auazAyord ay} Jo sureIDOIp] 


[ 221 ] 


TABLE II 
Chromosome lengths*, arm ratios, and percent relative contribution of each chromosome to the total genome length 
at pachytene in three species of Coix, viz., C. aquatica Roxb., C. Lachryma-Jobi L., and C. gigantea. 


Chromosome No. Species 
C. aquatica C. Lachryma-Jobi C. gigantea 
BS Percent Percent - Percent _ 
contribution contribution contribution 
Arm to the total Arm to the total Arm to the total 
Length ratio genomelength Length ratio genome length Length ratio genome length 

1 107.7 1.04 26.80 92.0 1.81 15.81 80.0 1.05 16.37 

m4 2 101.3 1.44 25.20 66.0 2.62 11.34 71.2 1.54 14.57 
S 3 84.6** 1.50 21.05 63.2 1.89 10.86 70.0 1.55 14.32 
in 4 56.9 2.33 14.16 59.2 4.58 10.17 67.2 1.70 13.75 
5 51.4 1.97 12.79 59.2 1.32 10.17 58.0 3.41 11.87 

6 52.0** 1.10 8.93 38.0 3.09 7.77 

7 50.0 2.08 8.59 37.6 2.07 7.69 

8 50.0 2.08 8.59 36.0** = 2.22 7.36 

9 45.6 1.04 7.84 30.8 1.39 6.30 

10 44.8 2.44 7.70 


*In microns. Mean value of up to five separate measurements in the case of C. aquatica and C. Lachryma-Jobi and ten 
measurements in the case of C. gigantea. 


** Nucleolus organizing chromosome. 


Jobi exhibits variation in chromosome morphology be- 
tween populations in two features: (a) the extent of 
differential staining into eu- and heteropycnotic regions ; 
and (b) the numbers and positions on chromosomes of 
knobs which are present on the chromosomes of this 
species. The chromosomes of C. gigantea show differen- 
tial staining as in the case of C. aquatica and some popu- 
lations of C. Lachryma-Jobi. But they are devoid of 
knobs. Comparative chromosome morphology does not 
reflect the presence of the common genome (or genomes) 
detected by interspecific hybridization. 

Comparison of chromosome morphology does not 
show similarities between C. aquatica, species of Sorghum 
with a diploid chromosome number of 10, and maize: 
whereas some similarities are evident between the chro- 
mosomes of C. Lachryma-Jobi and those of maize. Data 
on genome analysis based on intergeneric hybridization 
between Coix, Sorghum, and maize are not available in 
order to interpret in genetic terms the morphological 
comparisons made in this study. 


[ 223 ] 


LITERATURE CITED 


Anderson, E. 1945. What is Zea Mays? A report of progress. Chron. 
Bot. 9: 88-92. 

Bor, N.L. 1960. Grasses of Burma, Ceylon, India and Pakistan. 
Pergamon Press, New York. 

Chaganti, R.S.K. 1965, Cytogenetic studies of Maize-Tripsacum hy- 
brids and their derivatives. Bussey Inst. Harvard Univ. Publ., 
pp. 1-938. 

Darlington, C.D. and E.K. Janaki Ammal. 1945. Chromosome At- 
las of Cultivated Plants. George Allen and Unwin, London. 

Garber, E.D. 1947. The pachytene chromosomes of Sorghum intrans. 
Journ. Hered. 38: 251-252. 

Longley, A.E. 1941. Chromosome morphology in maize and its rela- 
tives. Bot. Rev. 7: 263-289. 

Mangelsdorf, P.C. and D.L. Oliver. 1951. Whence came maize to 
Asia? Bot. Mus. Leafl. Harvard Univ. 14: 263-291. 

Mangelsdorf, P.C. and R.G. Reeves. 1939. The origin of Indian 
Corn and Its Relatives. Texas Agric. Expt. Sta. Bull. No. 574: 
1-315. 

Mimeur, G. 1951. Systématique specifique du génre Coir et systé- 
matique varietale de Coir lacryma-jobi.. Morphologie de cette petite 
céréale et étude de sa plantule. Rev. Int. Bot. Appl. Agric. Trop. 
81: 197-211. 

Nirodi, N. 1955. Studies on Asiatic relatives of maize. Ann. Mo. 
Bot. Gard. 42: 103-130. 

Rees, H. 1961. Genotypic control of chromosome form and behavior. 
Bot. Rev. 27: 288-318. 

Reddi, V.R. 1958. On the differentiation of A and B chromosomes 
of Sorghum purpureo-sericeum at pachytene. Jour. Ind. Bot. Soc. 
387: 279-289. 

Rhoades, M.M. 1955. The cytogenetics of maize. In Corn and Corn 
Improvement. Ed. G.F. Sprague. Academic Press, New York, 
pp. 123-219. 

Venkateswarlu, J. 1958. Cytological observations on spontaneously 
occurring ring and chain formation in Cotx aquatica. Jour. Ind. 
Bot. Soc. 37: 329-333. 

Venkateswarlu, J. 1963. Cytogenetic evolution in angiosperms- 
Maydeae. Mem. Indian Bot. Soc. No. 4: 65-73. 

Venkateswarlu, J. and R.S.K. Chaganti. 1973. Job’s Tears. Tech. 
Bull. No. 44 Indian Council of Agricultural Research, New Delhi, 
pp. 1-54. 


[ 224 J 


NEW CHIHUAHUAN UMBELLIFERAE 
BY 
LINcoLN ConsTaNCE' AND Ropertr A. Byes, Jr.’ 


Our knowledge of the Umbelliferae of the Mexican 
Kstado de Chihuahua began apparently with Wislizenus’ 
pioneer journey to northern Mexico just before the mid- 
dle of the 19th Century, which yielded Hryngium hetero- 
phyllum Kngelm. The activities of the 1880s’ of those 
two doughty American collectors, Palmer and Pringle, 
led to the description of Arracacia edulis S. Wats., 

vryngium madrense S. Wats., Hulophus tenuifolius S. 
Wats., LZ. ternatus S. Wats., Museniopsis ternata var. 
Ailifolia C. & R., Prionosciadium madrense 8. Wats., and 
P. Pringle: S. Wats. A spate of field investigations at 
the turn of the century by Goldman, Nelson, M. E. 
Jones, and ‘Townsend and Barber led to the discovery 
of Contoselinum mexicanum C. & R., Eryngium fluitans 
M.E. Jones, 7. Goldmanii Hemsl., 7. medium Hemsl., 
Ligusticum Goldmani C. & R., L. madrense C. & R., 
L. Nelsonu C. & R., Museniopsis pubescens C. & R., and 
Prionosciadium Townsendi C. & R. Inthe 1980s’ there 
was another wave of collecting by Gentry, LeSueur, 
Muller and Pennell. Gentry’s intensive study of Rio 
Mayo plants led to the description of Hryngium calaster 
Standl. Up to the present, some thirty taxa of Umbel- 


' Department of Botany, University of California, Berkeley, Cal. 
* Botanical Museum, Harvard University, Cambridge, Mass. 


225 ] 


liferae are known to occur in the state, according to the 
taxonomic concepts of Mathias and Constance (1944- 
45). Since more than one-third of these taxa belong to 
the difficult genus Mryngium, this total is susceptible to 
change. 

In his field studies’ of the ethnobotany of the Tara- 
humara centered in the Sierra Madre of southwestern 
Chihuahua, the junior author has given particular atten- 
tion to Umbelliferae. As a result, he obtained material 
of no fewer than twenty taxa (including cytological ma- 
terial of many of them). Four of the collections repre- 
sent apparently undescribed species. The purpose of this 
paper is to put them adequately on record. 


Donnellsmithia silvicola Constance & Bye sp. nov. 

Plantae perennes caulescentes humiles condensatae ; 
caules simplices vel pauciramosae sparsim foliati 8-25 
em. alti e radice palari tumida omnino glabrae; folia 
basalia deltoideo-orbiculata 1-8 cm. longa, 1.5-5 cm. 
lata, trifoliata, foliolis 8 ovalibus orbiculatisve, obtusis 
lobatis vel pinnatifidis diametro 1-2 cm.: petioli 1-8 
cm. longi anguste scarioso-vaginantes; folia caulina ea 
basalia simulantia sed sursum reducta vaginis parum dila- 
tatis: pedunculi usque ad 2 em. longi pauci alternati 
graciles umbellis plerumque lateralibus sessilibus sub- 
sessilibusve; involucrum nullum:; involucellum nullum; 
radii 2—5 filiformes patenti-adscendentes 6-25 mm. longi: 
pedicelli fertiles 0-2, 3-4 mm. longi; flores flavo-virentes ; 
stylopodium depressum, stylis brevissimis ; carpophorum 
usque ad basim bipartitum; fructus immaturus ovoideo- 
orbicularis 1.5 mm. longus, 2 mm. latus ad apicem ver- 


* Financial support between 1971 and 1974 was generously extended 
by the Botanical Museum and the Department of Biology, Harvard 
University, the National Geographic Society and the National Science 
Foundation Grant GB-35047 for Improvement of Doctoral Disserta- 
tion Research. 


[ 226 | 


Tears LA 


a 


Donnellsmithia silvicola Const. & Bye. a, Habit, <4; b, basal leaf, 
“2; ¢c, flowering umbellet, «4; d, immature fruit, “10; e, petal, 
10. All from Bye 6376. 


sus truncatus basi rotundatus, costis humilibus filiformi- 
bus; vittae et seminum superficies non visae: chromoso- 
matum numerus 7 = 20. 

Small, compact perennials from a fleshy taproot, 8-25 
em. high, the stem simple or few-branched, very sparsely 
leafy, glabrous throughout; basal leaves deltoid-orbicu- 
lar, 1-8 em. long, 1.5—5 cm. broad, trifoliate with 8 oval 
to orbicular, obtuse, lobed to pinnatifid leaflets 1-2 cm. 
in diameter; petiole 1-8 cm. long, narrowly scarious- 
sheathing: cauline leaves like the basal but reduced up- 
ward, the sheaths slightly dilated; peduncles few, alter- 
nate, slender, up toe 2 cm. long, but most umbels lateral 
and sessile or subsessile; involucre 0, or of a single trifid 
bract; rays 2-5, filiform, spreading-ascending, 6-25 mm. 
long; involucel 0; fertile pedicels 0-2 (many umbellets 
staminate), 8-4 mm. long; flowers greenish-yellow; sty- 
lopodium depressed, not evident: styles very short; 
sarpophore 2-cleft to base; immature fruit ovoid-orbicu- 
lar, 1.5 mm. long, 2 mm. broad, truncate at apex, 
rounded at base, the ribs low, filiform: vittae and seed 
face not seen; chromosome number r= 20. 

Type. Chihuahua: Municipio de Guazapares, among Pinus ponderosa, 
P. leiophylla, Quercus sp. and Arbutus sp., on recently logged ridge, 
elevation 8100 feet, off road northwest of Estaci6n San Rafael to Las 
Lagunitas, 21 July 1974, Robert A. Bye, Jr. 6376 (GH: holotype; 
ECON, UC). 

The plant was rather abundant but very local on slopes 
that had recently been disturbed by logging operations. 

Donnellsmithia silvicola appears to be most closely re- 
lated to D. serrata (C. & R.) Math. & Const. and D. 
dissecta (C. & R.) Math. & Const., both species of south- 
ern Mexico. It is unlike both in its lower stature and 
glabrous foliage. From the former, it differs also in its 
shorter rays; from the latter, in its lack of an involucel. 
The most striking disparity, however, is in its uniquely 
distinctive leaf form. 

[ 228 | 


Tauschia bicolor Constance & Bye sp. nov. 

Plantae brevi-caulescentes acaulescentesve glabrae, 
‘caulibus gracilibus 10-40 cm. altis e caudice gracili: folia 
basalia rosulata triangulari-ovata vel ovata 1.5-8 cm. 
longa, 2-6 cm. lata, profunde trilobata vel ternata vel 
imparipinnata foliolis 1—8-partibus late ovalibus vel an- 
guste oblongis serratis lobatisque vel pinnatifidis, superne 
cinereis, inferne purpureis; petioli anguste vaginantes 
1.5-10(-25) em. longi; folia caulina pauca; pedunculi 
graciles 7-20 cm. longi; involucrum plerumque nullum: 
radii 8-12 eis exterioribus fertilibus 5-11 mm. longis: 
involucellum dimidiatum bracteolis 8-5 linearibus acutis 
1-3 mm. longis; pedicelli crassi 1-2 mm. longi fructo 
centrali saepe subsesseli; flores purpurei, petalis intus 
albis extus purpureis; stylopodium nullum, stylis valde 
gracilibus; carpophorum bipartitum dimidiis_ bifidis; 
fructus ovoideus apice truncatus basi rotundatus, glaber, 
2.5-8 mm. longus, 2 mm. latus, costis filiformibus acutis 
quam intervallis multo angustioribus; vittae in interval- 
lis plures in commissuris 2; seminum superficies plana; 
chromosomatum numerus 7 = 22. 

Plants rather slender, short-caulescent to acaulescent, 
the stems slender, 1 or 2, 10-40 cm. high from a slender 
rootstock, glabrous; basal leaves rosulate, triangular- 
ovate to ovate, 1.5-8 cm. long, 2-6 cm. broad, deeply 
3-lobed or ternate to pinnate with 1-8 pairs of broadly 
oval to narrowly oblong, serrate and lobed to pinnatifid 
leaflets 8-30 mm. long, 8-30 mm. broad, ashy-pale above, 
purplish beneath; petioles 1.5-10(—25) em. long, nar- 
rowly sheathing; cauline leaves few, reduced upward: 
peduncles terminal, slender, 7-20 cm. long: involucre 
of a linear bract or usually 0; rays 8-12, 5-11 mm. long, 
apparently only the outer fertile: involucel dimidiate, 
the bractlets 3—5, linear, acute, 1-8 mm. long; pedicels 
stout, 1-2 mm. long, the central fruit often subsessile ; 


[ 229 ] 


flowers purple, the petals white within, deep purple on 
the back: styles very slender, 1.5-2 mm. long, spread- 
ing, purple (contrasting with a white disc); carpophore 
2-parted, the halves bifid; fruit ovoid, 2.5-8 mm. long, 
” mm. broad, truncate at apex, rounded at base, the ribs 
filiform, acute, much narrower than the intervals; vittae 
several in the intervals, 2 on the commissure; seed face 


plane; chromosome number n= 22. 


Type: Chihuahua: near boundary of Municipio de Batopilas and 
Municipio de Urique, between Quirare and Basigochic, elevation about 
6900 feet, flowers red, 8 June 1973, Robert A. Bye, Jr. 4068 (GH: 
holotype; ECON, UC). It was collected in the same locality on 31 
May 1973, Bye 3863 (ECON, UC), and 6 October 1975, Bye 6965 
(ECON, UC). 


Tauschia bicolor, known to the Tarahumara as ‘‘masii- 
wari” (or ‘‘masiéwari’’)‘, is considered a quelite or pot- 
herb, although it is not commonly consumed today. The 
young tender leaves can be collected during the fall as 
well as during the spring. ‘To date, we know of only one 
locality for this herb, which is eagerly devoured by graz- 
ing goats and sheep. The plants are restricted to the 
western slopes above Arroyo Basigochic near an isolated 
stand of Abies durangensis. ‘They are common on moist 
(not wet) eroded slopes of partially open pine and oak 
forest, where they often grow with Chimaphila and 
Galium, while scattered individuals can be found in 
crevices of white volcanic rock or in thin soil of the mixed 
wood forest. Due to heavy grazing pressure, few plants 
are able to produce mature fruits. The rhizomes appear 
to be important in the vegetative propagation of the 


Pirate LXIL. Tauschia bicolor Const. & Bye. a, Habit, » 4: b, fruiting 
umbel, “2; c, mature fruit, “8; d, fruit transection, “15; e, carpo- 
phore, <8; f, petal, 15; g, first basal leaf, <4. a-d, f, from Bye 
4068; e, from Bye 6965; g, from Bye 3868. 


Ff a F ee x as 
* One informant referred to this herb as ‘“kurisove’’. 


[ 230 ] 


Puiate LXII 


population. Indeed, this characteristic may be the key 
to the survival of the population in the immediate future, 
because a road building program will destroy the site 
within the next year. 

Although plants relatively similar to T'auschia bicolor 
in habit may be found in such genera as -fletes and Arra- 
eacia as Well asin Tauschia, the combination of glabrous 
herbage, ternate or pinnate leaves with serrate to pinnati- 
fid leaflets, fertile outer rays, dimidiate involucel, stout 
pedicels with an often subsessile central fruit, petals 
purple/red externally and white internally, very slender 
styles, and a bipartite carpophore with bifid halves ap- 
pears to be unique. 


Tauschia tarahumara (Constance & Bye sp. nov. 

Plantae humiles breviter caulescentes glabrae, caulibus 
pluribus 12-30 cm. altis e caudice elongata ramosa car- 
nosa, praeter inflorescentiam puberulentam; folia ovata 
3-5 cm. longa, 2-4 em. lata, 2~-8-pinnata, divisionibus 
ultimis lineari-filiformibus breviter mucronatis 5-20 mm. 
longis, ca. 0.5 mm. latis; petioli 1-6 cm. longi graciles 
anguste scarioso-vaginantes; folia caulina reducta sessilia 
vaginis conspicue scariosis praedita; pedunculi 6-12 cm. 
longi plerumque terminales graciles sub umbella puberu- 
lentes; involucrum nullum; involucellum nullum ; radii 
6-12 usque ad 1 cm. longi puberulenti; pedicelli 1-38 
mm. longi eis exterioribus fertilibus: flores rhodo- 
purpurei; styli erecti ca. 0.5 mm. longi; carpophorum 
bifidum; fructus ovoideus 5-7 mm. longus, 8-5 mm. 
latus ad apicem versus truncatus basi rotundatus, costis 
humilibus rotundatisque quam intervallis latioribus; vit- 


Prare LXIITL. Vauschia tarahumara Const. & Bye. a, Habit, <4; b, 
basal leaf, “1; c, seedling, ~1; d, carpophore, 5; e, fruit tran- 
section, “7; f, mature fruit, » 4: g, flowering umbellet, 5; h, 
petal, ~ 20, a-b, g-h, from Bye 6288; c, from Bye 7027 (Const. 
1918); d-f, from 7027. 


PiatreE LXIII 


tae magnae in intervallis una in commisuris 2; seminum 
superficiesanguste suleata; chrosomatum numerus7= 44. 

Plants low, shortly caulescent, the stems several, 12- 
30 cm. high from a very elongate, branched, fleshy cau- 
dex, glabrous except for the puberulent inflorescence : 
leaves ovate, 8-5 cm. long, 2-4 cm. broad, 2—8-pinnate, 
the ultimate divisions linear-filiform, 5-20 mm. long, 
about 0.5 mm. broad, shortly mucronate; petioles slen- 
der, 1-6 cm. long, narrowly scarious-sheathing, the cau- 
line leaves with conspicuous scarious sheaths and sessile 
reduced blades; peduncles mostly terminal, slender, 6— 
12 cm. long, puberulent beneath the umbel: involucre 
and involucel 0; rays 6-12, up to 1 cm. long, puberulent ; 
pedicels 1-8 mm. long, the outer fertile; flowers deep 
red-purple; styles about 0.5 mm. long, mostly erect: 
carpophore bifid: fruit ovoid, 5-7 mm. long, 8-5 mm. 
broad, truncate at apex, rounded at base, the ribs low 
and rounded, broader than intervals; vittae large, soli- 
tary in the intervals, 2 on the commissure; seed face 
narrowly sulcate: chromosome number 7 = 44. 

Tyre: Chihuahua: Municipio de Bocoyna, in wettest portions of 
grazed meadow with dark mucky soil, elevation ca 7300 feet, N of 
San Ignacio Arareco, EF. of Creel, 11 July 1974, Robert 4. Bye, Jr. 
6288 (GH: holotype; ECON, UC). It was collected also in similar 
habitats in the general vicinity of San Ignacio Arareco, SE or E of 
Creel, on 14 July 1971, Bye 1535 (ECON), 16 July 1972, Bye 2369 
(ECON), 5 July 1973, Bye 4150 (GH, UC), and 19 July 1973, Bye 
4272B (ECON, UC), 9 October 1975, Bye 7027 (UC) and 7034 
(ECON). 


The older Tarahumara of San Ignacio call this herb 
“*huve™” and ascribe medicinal properties to the distine- 
tive rhizome. It is ground, mixed with oil or fat, and 
rubbed on the affected parts of the body to treat rheuma- 
tism. Only a small piece of the rhizome is placed in a 
cavity in order to alleviate toothache, because it is said 
to be very strong. Some young Tarahumara in the San 


| 284 | 


Ignacio region claim it is an edible green and call it 
“sene 

This deep rooted herb is restricted to a narrow inter- 
zone in the moist upland meadows, between the low, 
wet area (often occupied by standing water during parts 
of the rainy season) with Plantago and Ranunculus spe- 
cies, and the slightly elevated and drier area with J'agetes 
lucida. Even though this umbel has a deep, vertical root- 
stock (over 50 cm. long with a relatively constant di- 
ameter) which may be well adapted to environmental 
changes, such human-related activities as grazing, culti- 
vation, gully and sheet erosion, draining and soil com- 
paction in these accessible meadows threaten to destroy 
the few known populations of this curious plant. 

In the published key to the species of Tauschia 
(Mathias & Constance, 1944, pp. 81-82), the new taxon 
would key to either 7". tenuwifolia (S. Wats.) Math. & 
Const., a little known Chihuahuan plant, or to 7°. mari- 
ana (S. Wats.) C. & R. ex Drude (= 7\. decumbens 
(Benth.) C. & R. ex Drude), which occurs from México 
to Michoacan. It differs from both, however, by its lack 
of an involucel and its smaller fruit with broad, obtuse 
costae. In addition, 7". tarahumara is unlike 7". tenui- 
folia in lacking a densely fibrous stem base and in its more 
numerous but shorter rays. From 7°. decumbens, 7. 
tarahumara may be distinguished by its lower stature, 
broader leaves with narrower divisions, and red-purple 
flowers, in addition to its geographic range. 


Eryngium Gentryi Constance & Bye sp. nov. 

Plantae perennes graciles caulescentes haud ramosi 
1.8-5 dm. altae ex caudice horizontali gracili; folia pauca 
alternata disticha lineari-lanceolata 8-20 cm. longa, 2-5 
mm. lata, transverse septata, margine serrulato inte- 
grove, venis parallelis, vaginis amplexicaulibus latitudi- 


[ 235 ] 


nem laminae aequantibus vel longioribus; folia caulina 
summa opposita lanceolati-acuminata: inflorescentia 
cymosa reducta; capitula 1-8 globosa pedunculata sub- 
‘caerulea diametro 5-10 mm. ; bracteae involucrales 8-12 
lanceolatae acutae integrae vel pauci-dentatae, inferne 
virides, superne argenteae, quam capitulum duplo longi- 
ores: bracteae florales lineari-acuminatae integrae fruc- 
tum multo excedentes; sepala ovata ca. 2 mm. longa; 
petala lineari-oblonga 1—1.25 mm. longa apice fimbriata; 
styh graciles 2.5-8 mm. longi quam calyces longiores; 
fructus (immaturus) ovoideus diametro ca. 2 mm., super- 
ficiebus mericarpiorum dense squamatis, squamis ovatis 
subaequalibus complanatis; chromosomatum numerus 
n=T. 

Plants slender caulescent perennials 1.8-5 dm. high 
from a slender rootstock, the stems solitary to several, 
weakly erect, unbranched below inflorescence: lower 
leaves few, alternate, distichous, linear-lanceolate, 3-20 
em. long, 2-5 mm. broad, remotely serrulate or entire, 
the venation parallel, transversely septate, amplexicaul- 
sheathing, the sheath as broad or broader than the blade ; 
uppermost cauline leaves opposite, lanceolate-acuminate ; 
inflorescence a reduced cyme of 1-38 pedunculate heads: 
heads globose, bluish, 5-10 mm. in diameter; involucral 
bracts 8-12, spreading-ascending, lanceolate, 8-25 mm. 
long, 2-5 mm. broad, acute, entire or with one or two 
pairs of short teeth, green beneath, silvery-white above, 
about twice as long as the head: bractlets linear-acumi- 
nate, entire, much longer than fruit; sepals ovate, about 
2 mm. long, mucronate; petals linear-oblong, 1—1.25 
mm. long, with a narrower fimbriate tip; styles slender, 


Prater LXIV. Eryngium Gentryi Const. & Bye. a, Habit, <4; b, 
cauline leaf, «1; c, floral bractlet, «10; d, involucral bracts, 2; 
e, immature head (two bracts removed), <2; f, lower, 10; g, petal, 
<20. All from Bye 4766. 


[ 236 ] 


PuateE LXIV 


2.5-8 mm. long, longer than sepals; fruit (immature ) 
ovoid, about 2 mm. long and broad, the squamae ovate, 
subequal, flattened, the dorsal faces densely squamose ; 
chromosome number ”=7. 

Tyree: Chihuahua: Municipio de Ocampo, transition zone, pines, 
meadow, infrequent, marginal to streams, elevation 7500 feet, Meme- 
lichi, Rio Mayo, 16 September 1936, H.S. Gentry 2770 (UC: holo- 
type: kK). 


Oruer coLLections: Chihuahua: Municipio de Madera, lake near 


Chuichupa, 23 August 1936, H. LeSueur 817 (UC); Chuichupa, 7000 
feet, 21 September 19038, /.h. Diehl s.n. (POM): Municipio de Bo- 
coyna, in moist meadow E of Gonogochi (on Continental Divide) E 
of Creel, associated with low, scrubby Quercus, grasses, Ranunculus, 
Cacalia and Tagetes lucida, elevation ca 7500 feet, 24 August 1973, 


Robert 4. Bye, Jr. 4766 (ECON, UC). 


Kryngium Gentryi represents a third member of Sec. 
XN XI. Madrensia Wolff, comprising Mexican species of 
wet habitats characterized by evidently transversely sep- 
tate leaves, bluish or purplish ovoid-globose to ovoid- 
cylindrical heads, and involucral bracts conspicuously 
silvery above (Wolff 1918). The other two species, /. 
madrense S. Wats. and 7. fluitans M.K. Jones (27. meai- 
eanum sensu Wolff, 7. Wolffit Mathias), are very similar 
in habit. ‘There is a resemblance also to Ht. phyteumae 
Delar.f. (Sect. NNIV. Stellata Wolff), but this pos- 
sesses well developed oblong to lanceolate, reticulately 
veined leaf lamina. In Sect. Madrensia, the leaves are 
usually bladeless or with a rudimentary lamina and not 
only the petiole but the entire leaf tends to be septate. 
The silvery upper surface of the involucral bracts sug- 
gests an affinity to both Sects. NXNIV. Stellata Wolff 
and NNV. Carliniformia Wolff. Wolff notes that /. 
phyteumae ‘‘bildet einen deutlichen Ubergang zu den 
Madrensia’” (Wolff 1913, 61: 186). From its two allies 
in Sect. Madrensia, 47. Gentry: differs in its possession 
of a rootstock, its fewer globose heads without a coma, 


[ 288 | 


and in its broader involucral bracts that much exceed 
the head. 

The type collection was associated by Standley with 
the unpublished herbarium name ‘‘/. juncifolium M.E. 
Jones’’, which Jones had applied to specimens of both 
I. flutans and 7. Gentryt. Gentry reported the collec- 
tion in his Rio Mayo flora (1942) as H. phyteumae, as 
erroneously determined by Mathias and Constance. The 
“H. juncifolium Jones” listed by LeSueur in his pioneer 
study of Chihuahuan vegetation (1945) and attributed 
to the Montane Forest, is also 47. Gentry. The invalid 
name, ‘2’. juncifohum M.E. Jones’’, is not to be con- 
fused with the South American /1. juncifolium (Urban) 
Math. & Const. 

It is with great pleasure that we name this species in 
honor of Dr. Howard Scott Gentry, who has made such 
significant contributions to our knowledge of the flora 
of Chihuahua and other parts of Mexico. 


SPECIMENS 


Duplicates of the Bye collections will be deposited in 
the Herbario Nacional del Instituto de Biologia, Uni- 
versidad Nacional Aut6noma de México (MEXU),. 


[ 239 ] 


LITERATURE CITED 

Gentry, H.S. 1942. Rio Mayo Plants. Carneg. Inst. Wash. Publ. 
527, 328 pp. 

LeSueur, H. 1945. The ecology of the vegetation of Chihuahua, 
Mexico, north of Parallel Twenty-eight. Univ. Texas Publ. 4521, 
92 pp. 

Mathias, M.E., and L. Constance. 1944-45. Umbelliferae: Fl. N. 
America 28B: 43-295. 


Wolff, H. 1913. Umbelliferae; in Engler, Das Pflanzenreich 61 
(4°"): 1-305, 


[ 240 | 


CHROMOSOME NUMBERS IN 
CHIHUAHUAN UMBELLIFERAE 
BY 
LINCOLN ConstaNCcE!, Tsan [anc CHUANG? 
AND Ropsertr A. Byer, Jr.® 


In his ethnobotanical studies of the ‘Tarahumara cul- 
ture in the Sierra Madre of southwestern Chihuahua, 
Robert A. Bye, Jr., made a concerted and very success- 
ful effort to obtain material of Umbelliferae for chromo- 
some counts. The results of these counts (made by 
Chuang) are shown in the accompanying table and Plate 
LXV, and some brief comments (by Constance) follow 


(Table LT and Plate LXV, a-k). 


Donnellsmithia C. & R. The report on D. silvicola is 
new: that of 7=20 for D. ternata disagrees with one of 
n=21 for the same taxon (Constance, Chuang and Bell, 
in press). Chromosome numbers in this genus are still 
confusing. Counts made on eight specific taxa have 
yielded the following haploid numbers: 11, 14, 20, 21, 
22, 40, 42, 44. Thus, both D. silvicola and D. ternata 
are probably tetraploid. 

' Department of Botany, University of California, Berkeley, Cal. 

* Department of Biological Sciences, Illinois State University, 


Normal, II]. 


3 . ae . . 
Botanical Museum, Harvard University, Cambridge, Mass. 


C241 | 


EXPLANATION OF THE ILLUSTRATION 


Karyotypes of Chihuahuan umbelliferae 

a, Donnellsmithia ternata (S.Wats.) Math. & Const., Diak., Bye 4393: 
b, D. silvicola Const. & Bye, Diak., Bye 6376; ¢e, Tauschia bicolor 
Const. & Bye, Diak., Bye 3863; d, 7. tarahumara Const. & Bye, MII, 
Bye 6288; e, drracacia edulis S,Wats., Diak., Bye 6753; f, Eryngium 
Jluitans M.E.. Jones, MI, Bye 4763; g, FE. Gentryi Const. & Bye, 
Diak., Bye 4766; h, FE. heterophyllum Engelm., Diak., Bye 4334; i, 
Ek. Lemmonii Coult. & Rose (EF. calaster Standl.), MI, Bye 4463; j, 
EF. Lemmonii Coult. & Rose (FE. calaster Standl.), MII, Bye 4809; k, 
KE, phyteumae Delar.t., MIL, Bye 5905 (Const. 1906). (Diak.—= Dia- 
kinesis; MI, MII-Metaphases I and II. All ea. 1050). 


[ 242 ] 


PLATE LXV 


op a 
o 2 eo eC 
Qha .S J 4 
g ool o? ‘ 
6 go hy 
tf i 
> al o% 
r) ®e@ oe & 
oe oh ie Se 
q ay seve 
@ 5% ° 
Ne 4° -* 
eo 
Q eof 
© - a? "6 
e %°% Sf 
fle whe Ff 6? 
é 4 
a 
Po oj 


Tauschia Schlecht. Both 7’ bicolor and T. tarahumara 
were previously uncounted. Five other species of the 
genus have revealed only n=11 and n=22, so the count 
of n=44 for 7. tarahumara, a probable octoploid, rep- 
resents a new polyploid level for the genus. 


Arracacia Baner. There is no previous chromosome 
count for A. edulis, a species which has been placed al- 
ternately under T'auschia. The complement of 72=20 is 
equally discordant in both genera. The fifteen specific 
taxa of Arracacia examined heretofore have all shown a 
haploid number of 22, except for one unexplained occur- 
rence each of the numbers 14 and 82. 


Ligusticum L. The count of n=11 for L. Porter, 
which has been made before, agrees with findings on 
some 16 species of this cireumboreal genus, all of which 
showed n=11, n =22, or 2=33, save for an anomalous 
n=12 and one n=27. 

Pseudocymopterus C. & R. The count of 2=11 for 
P. montanus coincides with earlier findings for this genus 


and species. 


Prionosciadium S$. Wats. Of eight taxa of this genus 
counted, six have n=22, two have n=21. A count of 
22 for P. madrense conforms to earlier information. 


n= 


gryngium L. Of the six counts reported here, only 
that for #. Gentryi, n=7, is new. From some 100 spe- 
cies that have been counted in this very distinctive cos- 
mopolitan genus, it seems clear that the basic number is 
n=8. However, there are descending aneuploid series 
extending to n=7, 2 =6, or even nm=5 in both hemi- 
spheres, sometimes in what appears to be the same spe- 


[ 245 | 


cies, as in £7. heterophyllum and EB. Lemmoni. Extensive 
polyploidy, particularly in the Americas, has arisen on 
the basic complement of 8 (7 =16, 24, 82, 40, 48), and 
smaller series based apparently on 7 in Eurasia. 


LITERATURE CITED 
Constance, L., IT. 1. Chuang and C.R. Bell. Chromosome numbers in 
Umbelliferae V. Amer. Jour. Bot., in press. 


[ 246 | 


TABLE I 


Chromosome numbers of Chihuahuan Umbelliferae. 


Name 


Voucher 
Specimen* 


Municipio and Locality 


Donnellsmithia 
1. ternata(S.Wats.) C. & R. 


2, stlvicola Const. & Bye 
Tauschia 


3. bicolor Const. & Bye 


4. tarahumara Const. & Bye 


Arracacia 
5. edulis S.Wats. 


Ligusticum 
6. Porteri C. & R. 


Pseudocymopterus 


7. montanus (A.Gray)C.& R. 


Prionoseiadium 


8. madrense 5. Wats. 
Eryngium 

9. fluitans M.E. Jones 

10. Gentryi Const. & Bye 

11. heterophyllum Engelm. 


12. Lemmonii C. & R. 


13. phyteumae Delar.f. 


44: 


20 


11 


Bye 4393 


Bye 6376 


Bye 3863 


Bye 6388 


Bye 6753 


Bye 6651 


Bye 6637 


Bye 6352 


Bye 4763 


Bye 4766 


Bye 4334 


Bye 4463 
Bye 4809 


Bye 5905 


Guachochi, Cusdrare 


Guazapares, San Rafael- 
Las Lagunitas 


Batopilas—Urique, between 
Quirare and Basigochic 


Bocoyna, San Ignacio 
Arareco 


Guachochi, Cusdrare 


Bocoyna, W of Creel 


Bocoyna, W of Creel 


Batopilas, S of Quirare 


Bocoyna, W of Gonogochi 
Bocoyna, E of Gonogochi 
Bocoyna, Gonogochi 
Batopilas, N of Quirare 


Bocoyna, Creel—San 
Ignacio Arareco 


Bocoyna, W of Creel 


* Voucher specimens are preserved in ECON, GH and UC. 


ECONOMIC BOTANY LIBRARY 
OF OAKES AMES 
HARVARD BOTANICAL MUSEUM 


BOTANICAL MUSEUM LEAFLETS 
HARVARD UNIVERSITY 


CamBripGe, MassacnuusetTts, JuNE 30, 1976 VoL. 24, No. 9 


THE GENUS OECEOCLADES LINDL. 
BY 
Lestige A. GARAY AND PETER TayLor* 


In 1832 while describing Oeceoclades, as a genus differ- 
ent from Angraecum, Lindley offered the following ob- 
servation: ‘‘The genus Angraecum. . . is known by its 
undivided lip, which is neither cucullate, nor articulate 
with the column; by its spreading perianthium, which 
never has the segments turned upwards as in Eulophia; 
by its long taper-jointed spur, which is rarely enlarged 
at the base; and finally, by all these characters being 
connected with coriaceous leaves that are never ribbed 
or plaited. Such being the definition of Angraecum, it 
will be apparent that, . . . such as our Angraecum macu- 
latum and a few of those of Du Petit Thouars, must be 
excluded; these form a genus nearly related to Eulophia, 
from which they are to be distinguished by their coria- 
ceous leaves, by the perianthium never being secund, and 
by the want of a crest upon the lip.” 

With this introduction Lindley also provided a sepa- 
rate generic description in Latin as well as a list of names 
—five under Angraecum and four under Limodorum — 
which he considered belonged to Oeceoclades, but with- 
out making the proper nomenclatorial transfers. ‘The 
following year, in 18338, in his Genera and Species of 
Orchidaceous Plants, he treated Oeceoclades in a similar 


* Orchid Herbarium, Royal Botanic Gardens, Kew, England. 


[ 249 | 


fashion, but with a somewhat different content. On both 
occasions, however, he included the sympodial Kulophia- 
like Angraecum maculatum with the remainder, all 
monopodial orchids. At that time Lindley remarked: 
‘‘Oeceoclades will probably comprehend all the Eulophia- 
like epiphytes... ; it is very near Eulophia, from which 
its coriaceous, not plaited leaves, distinguish it among 
other things. ”’ 

Thus, from the very beginning Lindley considered 
Oecceoclades to represent Eulophia-like plants. He em- 
phasized his conviction once more in 1859 (Journ. Linn. 
Soc. 83: 86) by saying ‘*... it is probably that O. macu- 
lata is the only plant to which the generic name will 
attach.’ Since that time the remaining species of Oeceo- 
clades have all been transferred to various angraecoid 
genera. Consequently the genus Oeceoclades must be 
typified by O. maculata, the only Eulophia-like plant. 

In 1887, Pfitzer in his Entwurf einer natiirlichen An- 
ordnung der Orchideen p. 87-88 established the mono- 
typic genus Mulophidium which he also based on Angrae- 
cum maculatum. It is difficult to understand why he chose 
to follow that particular course when he was fully aware 
of Lindley’s suggestion that the name Oeceoclades be 
applied only to O. maculata: ‘‘Ich muss Hulophidiwm 
maculatum entsprechend Lindley’s Vermuthung als den 
Typus einer besondren Gattung betrachten. . . “— 

Since both genera, Oeceoclades Lindl. and Hulophidium 
Pfitz. are based on the same type—Angraecum macula- 
tum V.indl.—Oececeoclades must be reinstated because of 
the rule of priority. 

Summerhayes, in 1957, published a synopsis of the 
genus Hulophidium (Bull. Jard. Bot. Bruxelles 27 (3): 
391-403). In that study he argued that Pfitzer and sub- 


** In keeping with Lindley’s supposition, I must regard Eulophidium 
maculatum as the type of a special genus.”’ 


[ 250 ] 


sequently Schlechter, both in their studies emphasized 
the vegetative aspects of the plants without paying much 
attention to floral details, which resulted in a rather poor 
circumscription of the genus. 

‘l'o augment this one-sided presentation, Summerhayes 
provides the following observations: 

‘*For some time now I have been struck by the marked 
similarity in floral structure between typical members of 
Eulophidium, such as F. maculatum (LINDL.) PFITZ., 
on the one hand, and species which have always been re- 
tained in Hulophia such as I. saundersiana RCHB.F. 
and the Asiatic 17. macrostachya LIN DL., on the other. 
All these species have a marked quadrilobed labellum 
with two short parallel or slightly divergent calli at the 
base and no long keels or hair-like outgrowths such as 
are so widely distributed in Mulophia. The side lobes 
almost invariably have marked darker veins. . . Some- 
time the two lower lobes, or lateral lobes if you prefer 
to call them that, are much reduced, occasionally so 
much that the labellum is almost bilobed. The spur is 
relatively short and often swollen, sometimes it is more 
or less shortly bilobed at the apex. 

‘*As regards the vegetative structure the aerial pseudo- 
bulbs may be heteroblastic with 1-8 leaves at the apex 
or homoblastic (with several elongated internodes) with 
one or more leaves at the apex. All intermediates can 
be found between a clearly heteroblastic condition with 
no cataphylls or leaves arising along the pseudobulbs, 
through forms in which the lower swollen internodes are 
quite short and other forms with only 2 elongated swol- 
len internodes, to typical homoblastic conditions where 
there are clearly several well-defined elongated swollen 
internodes with cataphylls arising from the lower nodes 
and leaves from the uppermost. 

“The great majority of species have markedly petio- 


[ 251 | 


late leaves, but in a few cases the petiole is very short. 
It is almost invariably articulate with a number of sharp 
or blunt teeth at the joint, this articulation usually being 
some distance above the base of the leaf, and sometimes 
in the centre of the long slender petiole. Reichenbach 
and Schlechter, when dealing with some species, have 
treated the lower part of the petiole below the articula- 
tion as being the rostrate upper internode or prolonga- 
tion of the pseudobulb, but this is clearly not a possible 
interpretation in plants like J”. sawndersiana where there 
are two leaves, both with their petioles articulate some 
distance above the base. The leaves are usually rather cori- 
aceous and often banded or spotted with paler markings. 

‘* As these variable characters are associated with very 
similar floral structures I feel that too much emphasis 
should not be placed on them and that all these species 
should be placed in an enlarged Hulophidium.” 

In no way diminishing the value of these important 
observations, we are compelled to reexamine some of its 
crucial points. We believe that the distinction between 
homoblastic and heteroblastic pseudobulbs is not clearly 
understood by many. Both types of pseudobulbs are de- 
rived from the common sympodial stem which consists 
of aset of nodes and internodes. In the case of the homo- 
blastic pseudobulbs, each internode or most of the inter- 
nodes are equally developed and enlarged throughout 
the entire length of the sympodium, at the nodes leaves 
or sheaths or cataphylls are produced. In the case of the 
heteroblastic pseudobulbs only one internode is fully de- 
veloped and enlarged throughout the entire length of the 
sympodium. The nodes are closely approximate, often 
so congested that no visible separation is apparent ; rarely 
the leaf-bearing nodes are separated by obvious distances, 
yet minimal in proportion to the internode that repre- 
sent the true heteroblastic pseudobulb. The one to three 


[ 252 | 


leaves produced at the top of a heteroblastic pseudobulb 
are originating independently from separate nodes, which 
may or may not be closely approximate. The misunder- 
standing of the nature of the heteroblastic pseudobulb 
caused Summerhayes to enlarge and alter the generic 
description of Mulophidium, which unfortunately now 
encompasses several criteria applicable only to Hu/lophia. 

Likewise the similarities in floral structure, mentioned 
by him, we believe are produced through convergent 
evolution rather than through the processes of speciation. 

Although florally Hulophia macrostachya is very simi- 
lar to those found in many plants of Oeceoclades, the 
thin, plicate leaves and the homoblastic pseudobulbs 
immediately exclude it from that relationship. If FH. 
macrostachya is to be admitted to Oeceoclades, then FE. 
graminea, I’. euglossa, E.. guineénsis, and other related 
species would have to be included also. For additional 
names see the list of Mulophia Sect. Pulchrae Kral. at 
the end of this paper. 

Both Lindley and Pfitzer were explicit about such 
generic characters as the heteroblastic pseudobulbs, cori- 
aceous, conduplicate leaves and Eulophia-like flowers. 
Consequently we adhere to the original circumscription 
of the genus in our assignment of the species. 


Oeceoclades Lindl. in Bot. Reg. 18: sub t. 1522, Sept. 
1, 18382. 
Syn.: Aeceoclades Duch. in Orbigny, Dict. 9: 170, 
1849. 
Saccolabium Sect. Oecceoclades (Lindl.) Cor- 
dem., Fl. Reunion 197, 1895. 
Lectotype: Angraecum maculatum  Lindl.— 
Lindl]. in Journ. Linn. Soc. 3: 36, 1859. 
Eulophidium Pfitz., Entw. Natur. Anordn. 
Orch. 87-89, 1887. 


[ 253 ] 


Eulophia Sect. Hulophidium (Pfitz.) H. Perr. 
in Bull. Soe. Bot. Fr. 82: 147, 19385. 
Lissochilus Subgen. Hulophidium (Pfitz.) H. 
Perr. in Humbert, Fl. Madag. Orch. 2: 17, 
1941. 

Type: Angraecum maculatum Lindl.—Pfitz. did. 


Sepals and petals variously spreading; lip 8-lobed, 
basally produced in a spur, midlobe commonly lobulate 
or emarginate; disc either with a pair of approximate, 
quadrate or triangular calli at the entrance to the spur 
or with three variously thickened, parallel ridges which 
together with the lateral veins are sparsely but distinctly 
papillose or hirsute; column erect, rather short, oblique 
at base; stigmata confluent; rostellum short; anther 
cucullate to cristate; pollinia 2, on a short or rudimen- 
tary stipe; viscidium large. 

Pseudobulbs more or less approximate, heteroblastic, 
one- to three-leaved at apex; leaves coriaceous, condu- 
plicate, never plicate, commonly petiolate, rarely sessile, 
articulate with colliferous apex of pseudobulbs; inflores- 
cence lateral, racemose or paniculate; bracts inconspicu- 
ous; flowers rather small and thin in texture, resupinate. 

31 species native to tropics and subtropics of Sey- 
chelles, Madagascar, the Mascarene Islands, Africa, 
South America, West Indies and Bahamas. 


Type of the genus: Angraecum maculatum Lindl. 


Key to Species 


1. Petals at most 4 to 4 the length of the sepals . . . . . . 2 
la. Petals and sepals more or less equal in length . . . . . . 6 
2, Pseudobulbs ovoid to cylindrical; leaves lorate [at 

least 40 em. long]; inflorescence diffusely branched. . . . 3 


2a. Pseudobulbs globose to pyriform; leaves linear [at 
most 20 em. long]; inflorescence racemose or rarely 
with few short branches . . . . . e 7 ee ee ee ee 4 


[ 254 ] 


3a, 


8a. 


9a, 


Pseudobulbs 2-leaved; sepals spathulate to oblan- 

ceolate, obtuse; petals elliptic, obtuse; spur of lip 

forward projecting under lip... .. . . .. . O.calcarata 
Pseudobulbs 1-leaved; sepals and petals lanceolate- 

elliptic, acute to subacuminate; spur of lip project- 


ing away fromlip. ......... .. .. O. Hebdingiana 
Sepals spathulate, obtuse; spur globose . . . O. spathulifera 
. Sepals obovate-oblanceolate, acute. . .......4.. 5 


Leaves sessile; sepals at least 14 mm. long; petals 

elliptic, acute; spurcylindric. . .... . . O. Decaryana 
Leaves petiolate; sepals not more than 8 mm. long; 

petals suborbicular, obtuse; spur subglobose, ven- 


trally compressed. . ... . 2. . « . OJ angustifolia 
Lip ecallose at base; basal Baise of 3 parallel veins 
of disc somewhat carinate-thickened . ... . - ee ee 


Lip with a bilobed callus, or bilamellate at or near Bee - » « 1S 
Pseudobulbs elongate, slender, fusiform to cylindric, 

2-3 leaved, approximately the length of the petiolate 
feaves or longer 4-5. we ew: eB a ae wwe me ee a 8 
Pseudobulbs short, ovoid, 2-leaved, much shorter 

than the petiolate leaves. . . A = 
Lip broadly elliptic; midlobe of lip senda: at oe 
overlapping with lateral lobes without a sinus; disc 

ecallose at junction of lateral and median lobes ..... 9 
Lip narrowly ovate-oblong to elliptic-oblong; midlobe 

of lip cuneate at base, forming a distinct sinus with 

lateral lobes; disc with a pair of fleshy gibbosities at 

junction of lateral and midlobes. . . . .. . . O.ugandae 
Midlobe of lip suborbicular in outline, half as long 

as the entire length of the lip; lateral lobes subfal- 

cate, obtuse at apex. .. . . « .  O, lanceata 
Midlobe of lip reniform in eee one- fourth the 

length of the entire lip; lateral lobes broadly 


rounded atapex ........... . . O.seychellarum 
Lip lobate from middle; lateral lobes of lip 
teunente in ont «6 6 «ow a 4: 4 ee ee ee oe DT 


Lip lobate one-fourth from apex; lateral lobes of lip 

OUUduE 1 TORE 6s a eG Se we RS De HS SB AB 
Lip wider than long; midlobe deeply emarginate to 
divaricately bilobulate in front . .. . . . .O. Lubbersiana 
Lip longer than wide; midlobe cuneate or with a 

distinct claw, at most retuse to indented in front O. pandurata 
Inflorescence laxly racemose; flowers greenish- 


[ 255 


16. 


16a. 


18a, 


19a. 


20. 


20a. 


21. 


yellow with maroon dots; sepals and petals 


ovate-oblong, acute... . ; . 2 2. « « O. atrovirens 

Inflorescence with short ecnelieds ncn white, 

sepals and petals linear-oblong, obtuse . . . . . O. latifolia 

Blades of leaves narrow, linear . . . . ..... =... 14 
a. Blades of leaves broad, ovate to elliptic. . . ..... .17 


Leaves long-acuminate, lorate, 50 cm. or more long, 
gradually tapering to pseudobulb without a distinct 

Wellgle 4. 9 40% % « & & &'% ce eae oo & OD Perse 
Leaves acute or obtuse, linear- sieve: 30 cm. or less 

long, petiolate or subpetiolate .. . a ¢ aa oe 
Spur vesicular, longer than lip; median aes of lip 

reflexed; disc in front of callus inormate . . . O. quadriloba 


. Spur cylindrical, shorter than lip; median lobe of 


lip not reflexed; disc in front of callus 3-carinate . . . . 16 
Pseudobulbs 2-leaved ; leaves subpetiolate, 20-30 

cm. long; inflorescence racemose; lateral lobes of 

lip rounded, larger than median lobe . . . . O. sclerophylla 
Pseudobulbs 1-leaved; leaves petiolate, less than 

15 cm. long; inflorescence branched ; lateral lobes 

of lip subquadrate with obtuse angles, equal to, or 

somewhat smaller than median lobe . . . . QO. analavelensis 
Plants caespitose ; pseudobulbs aggregate, ecol- 

liferous or with hardly any projections; leaves 

sessile or with conduplicate, short, petiole-like 


base ... ‘ ee eo hie 2 ow ee ole 
Plants diicsdutouxt : spendouaihe ners. 

prominently colliferous, leaves distinctly petiolate . . . . 25 
Pseudobulbs 2-leaved . ......... ++ 4. 419 
Pseudobulbs l-leaved . . . . 1... 1 ee ee ee ee 20 


Inflorescence profusely paniculate; lateral lobes of 

lip faleate, when expanded parallel with, and as long 

as midlobe; midlobe of lip wider than long; spur 

conical, acuminate ..... . 2. ee «OJ gracillima 
Inflorescence racemose or rarely with one or few 

short branches; lateral lobes of lip triangular to 

oblong, when expanded rectangular with midlobe ; 

midlobe of lip as wide as long, subquadrate; spur 


vesicular, obtuse. . . . .-. . . +. . . QO. roseovariegata 
Leaves ovate, subcordate at base... ....... + 21 
Leaves elliptic to oblong-elliptic, cuneate at base . . . . 22 


Sepals and petals similar, elliptic to ovate-elliptic, 
obtuse; lateral lobes of lip much larger than mid- 


[ 256 | 


Qla. 


22. 


22a, 


28a. 


29a. 


30. 


30a. 


lobe; column short, erect . . . .. . . . . . O. boinensis 
Sepals and petals dissimilar, linear-lanceolate, 

acute; lateral lobes of lip as large as midlobe; 

column elongate, arcuate ......... . . O. Rauhii 
Lip as long as or shorter than wide; terminal lobe 

separated from the lateral lobes by acute, indented 

or rounded sinuses .... . a a ae ee ee 
Lip as long as or longer than wide: terminal lobe 

separated from the lateral lobes by a distinct 


isthmus... . . . « O. monophylla 
Spur eapelonece: ee of lip bilobed . . . . O. ambongensis 
. Spur clavate to cylindric; callus of lip bilamellate . . . . 24 


Sepals and petals acute: lip as long as wide; sinuses 

in middle of lip; inflorescence always racemose . O. maculata 
Sepals and petals obtuse; lip shorter than wide; 

sinuses one-third from apex of lip; inflorescence 

racemose or subpaniculate .. . » . . « « OF Mackeni 
Pseudobulbs narrowly cylindrical, often stem-like, 

terete, two-leaved; lateral sepals longer than dor- 


sal sepal; lip equally 4-lobed . . . . . . . O. Saundersiana 
Pseudobulbs ovoid to conical, 1-leaved; sepals of 

same length; lip flabellate or unequally 4-lobed . . . . . 26 
Lateral lobes of lip larger than median lobe. . . . . . . 27 
Lateral lobes of lip much smaller than median 

lobe, ear-like . ... . & ; : ; ee 4 ee 
Leaves lanceolate, joni-abeenuaie, si bachametes 

sepals and petals apiculate . . . fu ea 4 0. sansibarica 


Leaves lanceolate-elliptic or Sacco elliptic, 

acute to subobtuse; sepals and petals obtuse O. alismatophylla 
Callus of lip and the disc in front of it puberulent . . . . 29 
Callus of lip and dise glabrous. . ‘5 - A moo 
Leaves broadly elliptic, large, 7- caened. with esti 

30-50 em. long; column-wing ciliolate-hirsute along 

margins; lateral lobes of lip rounded. . .  O. cordylinophylla 
Leaves narrowly ovate-lanceolate, 1-3 nerved, with 

petiole 13-20 cm. long; column-wings glabrous along 
margins; lateral lobes of lip obliquely triangular- 

faleate ... . 2. 2. . O. analamerensis 
Lip donuheleted 4 in aaidale. sccicniate: basal part 

suborbicular, apical part divergingly bilobed with 

rounded lobes; disc bilamellate at base in front 

of which are 8 thickened veins . ..... . . O. pettolata 
Lip not pandurate, 3-lobed; lateral lobes obtusely 


[ 257 | 


angular, semiovate, terminal lobe divergingly 

bilobulate with rectangular-oblong lobules; 

disc without thickened veins in front of basal 

lamellae . ........... . . . . 0. lonchophylla 


ENUMERATION OF SPECIES 


Oeceoclades alismatophylla (Rchb.f.) Garay & 
Taylor, comb. nov. 
Basionym: HMulophia alismatophylla Rehb.f. in Flora 
68: 5438, 1885. 
Syn.: Mulophidium alismatophyllum (Rehb.f.) Sum- 
merh. in Bull. Jard. Bot. Bruxelles 27: 894, 
1957. 
Type: Madagascar, Forét d’Ankaye. Coll. Humblot 
s.n.f (W). 
Distribution: Madagascar. 


Vegetatively the plants of this species are very similar to O. anala- 
merensis, O. petiolata and O. lonchophylla, but readily distinguishable 
from them in the floral structure, especially in the shape of the lip 
with the lateral lobes being larger than the median Jobe. We have 
seen only the type specimen. 


Oeceoclades ambongensis (Schltr.) Garay & Tay- 
lor, comb. nov. 
Basionym: HMulophidium ambongense Schltr. in Ann. 
Mus. Col. Marseille, ser. 3, 1: 182, t. 17, 
1913. 
Syn.: Hulophia Schlechtert H. Perr. in Bull. Soc. Bot. 
Fr. 82: 154, 1935. 
Lissochilus Schlechteri (H. Perr.) H. Perr. in 
Humbert, Fl. Madag. Orch. 2: 27, 1941. 
Type: Madagascar, Manongarivo (Ambongo). Coll. 
Perrier no. 1684! (P). 
Distribution: Madagascar. 


From the related species of the O. maculata alliance, the plants of 
this species are readily identifiable by the comparatively larger flow- 
ers, subglobose spur and the bilobed callus of the lip. 


[ 258 ] 


Oeceoclades analamerensis (H. Perr.) Garay & 
Taylor, comb. nov. 
Basionym: Lissochilus analamerensis H. Perr. in Not. 
Syst. 8: 42, 1939. 
Syn. : Hulophidium analamerense (H. Perr.) Summerh. 
in Bull. Jard. Bot. Bruxelles 27: 894, 1957. 
Type: Madagascar, Province de Diégo-Suarez, Anal- 
amera, rive droite de la riviére Analabe, afflu- 
ent du Rodo. Coll. Humbert no. 19247! (P, Kk). 
Distribution: Madagascar. 


The two small approximate lamellae near the base of the lip and 
the hirsute dise of the lip amply separate the plants of this species 
from the vegetatively similar O. alismatophylla. 

Perrier gives Humbert no. 19020 as the type number, but the speci- 
mens in Paris as well as at Kew bear the number 19247! 


Oeceoclades analavelensis (H. Perr.) Garay & Tay- 
lor, comb. nov. 
Basionym: Lissochilus analavelensis H. Perr. in Not. 
Syst. 8: 41, 1939. 
Syn.: Hulophidium analavelense (H. Perr.) Summerh. 
in Bull. Jard. Bot. Bruxelles 27: 895, 1957. 
Type: Madagascar, Forét d’Analavelona, au N. du 
Fiherenana. Coll. Humbert no. 14218! (P,K). 
Distribution: Madagascar. 


Florally the plants of this species are very similar to those of O. 
sclerophylla, both having three thickened ridges in front of the callus, 
but they are very dissimilar vegetatively. 


Oeceoclades angustifolia (Sengh.) Garay & Taylor, 
comb. nov. 
Basionym: Hulophidium angustifolium Sengh. in 
Adansonia ser. 2, 6: 558, 1967. 

Type: Madagascar: near Diégo-Suarez. Coll. Rauh 
& Buchloch no. 7987 (HEID). 

Syn.: Hulophidium angustifolium ssp. diphyllum 
Sengh. in Adansonia ser. 2, 6: 561, 1967. 


[ 259 | 


Type: Madagascar, near Sakaraha, river Fiherenana. 
Coll. Rauh no. 10423 (HEID). 

Distribution: Madagascar. 

From the related O. Decaryana, the plants of this species are dis- 


tinguished in having petiolate leaves as well as differently propor- 
tioned lips. 


Oeceoclades atrovirens (Lindl.) Garay & Taylor, 
comb. nov. 
Basionym: HMulophia atrovirens Lindl., Gen. and Sp. 
Orch. Pl. 184, 1833. 

Syn.: Graphorchis atrovirens (Lindl.) O. Ktze., Rev. 
Gen. Pl. 2: 662, 1891. 

Type: India, without proper locality. Coll. Wallich 
s.n.! (KX). 

Distribution: India orientalis. 


This peculiar plant is known from a colored drawing prepared by 
Wallich in 1828 for the East India Co., now in the Kew Herbarium. 
Ironically, Lindley originally bas written on the drawing ‘‘Oeceo- 
clades’’, then at a later time he crossed it out. So far no specimens 
are known to exist of this plant. Judging from the drawing, especially 
the floral details, it is near O. latifolia from which it differs in having 
a simple raceme, rather pointed sepals and petals and a somewhat 
different lip. 


Oeceoclades boinensis (Schlitr.) Garay & Taylor, 
comb, nov. 
Basionym: Mulophidium boinense Schltr. in’ Ann. 
Mus. Col. Marseille ser. 8, 1: 182, t. 17, 
19138. 
Syn. : Lissochilus boinensis (Schlitr.) H. Perr. in Hum- 
bert, Fl. Madag. Orch. 2: 26, 1941. 
Type: Madagascar, bords de la riviére Andranofasy 
(Boina). Coll. Perrier no. 1884! (P). 
Distribution: Madagascar. 
This species has its only relative in O. Rauhi, both having a more 
or less cordate base to the leaves. In floral structures they are, how- 


ever, very different. The type number of O. boinensis is Perrier 1834! 
and not 1384 as given by Schlechter. 


' 260 ] 


Oeceoclades calcarata (Schltr.) Garay & Taylor, 
comb. nov. 
Basionym: Cymbidium calcaratum Schltr. in Ann. 
Mus. Col. Marseille ser. 8, 1: 181, t. 16, 
19153. 
Syn.: Hulophia calcarata (Schltr.) Schltr. in Fedde 
Rep. Beih. 32: 262, 1925. 
Type: Madagascar, Manongarivo (Ambongo). Coll. 
Perrier no. 1681! (P). 
Eulophia paniculata Rolfe in Gard. Chron. ser. 
3, 88: 197, 1905, not O8cceoclades paniculata 
Lindl. 
Lissochilus paniculatus (Rolfe) H. Perr. in 
Humbert, Fl. Madag. Orch. 2: 29, 1941. 
Eulophidium paniculatum (Rolfe) Summerh. 
in Bull. Jard. Bot. Bruxelles 27: 399, 1957. 
Type: Madagascar, without precise locality. Collec- 
tor unknown. Flowered in cultivation at the 
Royal Botanic Garden, Glasnevin in June 
1904! (K). 
Distribution: Madagascar. 


The forward-projecting spur under the lip is unique in the genus. 


Oeceoclades cordylinophylla (Rchb.f.) Garay & 

Taylor, comb. nov. 

Basionym: Eulophia cordylinophylla Rehb.f. in Flora 

68: 541, 1885. 

Syn.: Lissochilus cordylinophyllus (Rehb.f.) H. Perr. 
in Humbert, Fl. Madag. Orch. 2: 20, 1941. 
Eulophidium cordylinophyllum (Rcehb.f.) Sum- 
merh. in Bull. Jard. Bot. Bruxelles 27: 3895, 
1957. 

Type: Comoro Islands, without proper locality. Coll. 
Humblot s.n. (W). 


Eulophia lokobensis H. Perr. in Bull. Soc. Bot. 
Fr. 82: 158, 1985. 


[ 261 } 


Lissochilus lokobensis (H. Perr.) H. Perr. in 
Humbert, Fl. Madag, Orch. 2: 22, 1941. 
Eulophidium lokobense (H. Perr.) Summerh. 
in Bull. Jard. Bot. Bruxelles 27: 8396, 1957. 
Type: Madagascar, Forét de Lokobe dans I|’Ile de 
Nossi-Bé. Coll. Perrier no. 19013! (P). 
Distribution: Comoro Islands, Madagascar. 


The actual specimen of Eulophia cordylinophylla is missing. How- 
ever, there is a sheet among Humblot’s collections from the Comoro 
Islands with an unpublished name by Reichenbach, which agrees well 
with the original description. This specimen also is identical with the 
type of FE. lokobensis from He de Nossi-Bé, near the Comoro chain. 
Perrier’s description of the column-wing as being toothed is incorrect. 
The margins of the column are ciliolate-hirsute. 


Oeceoclades Decaryana (H. Perr.) Garay & Taylor, 
comb, nov. 

Basionym: Hulophia Decaryana H. Perr. in Bull. Soe. 

Bot. Fr. 82: 154, 1935. 

Syn.: Lissochilus Decaryanus (H. Perr.) H. Perr. in 
Humbert, Fl. Madag. Orch. 2: 32, 1941. 
Vulophidium Decaryanum(H.Perr.)Summerh. 
in Bull. Jard. Bot. Bruxelles 27: 395, 1957. 

Type: Madagascar, without proper locality. Coll. 

Decary s.n. (P). Type is cultivated in Serres 
du Museum d’Histoire Naturelle de Paris 

(no. K 467). 
Distribution: Madagascar, Mozambique, Rhodesia, 

Kenya. 

Judging from the assortment of specimens we have examined, the 
length of the sepals varies with age. It may be separated easily from 
the related O. spathulifera by the shape of the spur. Because of the 


cylindrical spur, the illustrations published in Die Orchidee 18: 246, 
1967, as Eulophidium spatuljferum are referable here. 


Oeceoclades gracillima (Schltr.) Garay & Taylor, 
comb, nov. 


[ 262 ] 


Basionym: Hulophia gracillima Schltr. in Ann. Mus. 
Col. Marseille ser. 3, 1: 170, t. 14, 1913, 
not Ridl. 1886. 
Syn.: Mulophidium gracillimum Schltr. in Fedde Rep. 
Beih. 88: 255, 1925, nom. nov. 

Lissochilus gracilimus (Schltr.) H. Perr. in 
Humbert, Fl. Madag. Orch. 2: 28, 1941. 
Type: Madagascar, bassin du Besafotra, affluent de 

Menavava (Boina). Coll. Perrier no. 1059! (P). 
Distribution: Madagascar. 


This species is closely related to O. roseovariegata, but the diffusely 
paniculate inflorescence and the shape of the lip and spur readily keep 
them apart. 


Oeceoclades Hebdingiana (Guillaum.) Garay & 
Taylor, comb. nov. 
Basionym: Lissochilus Hebdingianus Guillaum. in 
Bull. Mus. Nat. Hist. Nat. ser. 2, 35: 
521, 1968. 
Type: Madagascar, sous bois d’Anipanihy, Prove- 
nance Montagnac. Flowered in cultivation in 
Jardin Botanique ‘‘ Les Cedres’’. Collector 
unknown! (P). 
Distribution: Madagascar. 


Related to O. calcarata from which it differs primarily in the shape 
of the lip and not having a forward-projecting spur. 


Oeceoclades lanceata (H. Perr.) Garay & Taylor, 
comb. nov. 
Basionym: Mulophia lanceata H. Perr. in Bull. Soe. 
Bot. Fr. 82: 156, 19385. 
Type: Madagascar, bois des pentes occidentales a 
Manerinerina sur le Tampoketsa, entre |’ [kopa 
et la Betsiboka. Coll. Perrier no. 168438! (P). 
Distribution: Madagascar. 


Perrier has reduced this species to a synonym of Eulophia pandu- 
rata Rolfe, but the two are amply distinct from one another in floral 


[ 263 ] 


details, especially in the shape of the lip. Vegetatively it is reminis- 
cent of O. seychellarum. The flowers are rose-colored. 


Oeceoclades latifolia (Rolfe) Garay & Taylor, comb. 
nov. 
Basionym: Mulophia latifolia Rolfe in Bol. Soc. Broter. 
9: 189, 1891. 
Syn.: Mulophidium latifolium (Rolfe) Summerh. in 
Bull. Jard. Bot. Bruxelles 27: 896, 1957. 
Type: Island of Sio Tomé. Coll. Quintas s.n./ (KX). 
Distribution: Africa—Island of Sao Tomé. 
Florally the plants referable to this species are rather similar to 
O. ugandae, but vegetatively they are very different and more remi- 
niscent of O. atrovirens. The lateral veins of the lip are papillose- 


ciliolate in this alliance to which O. pandurata, O. seychellarum and 
O. lanceata also belong. 


Oeceoclades lonchophylla (Rchb.f.) Garay & Tay- 
lor, comb. nov. 
Basionym: IZulophia lonchophylla Rchb.f. in Flora 68 : 
542, 1885. 
Syn.: HMulophidium lonchophyllum (Rehb.f.) Schltr. in 
Fedde Rep. Beih. 83: 256, 1925. 
Lissochilus lonchophyllus (Rehb.f.) H. Perr. in 
Humbert, Fl. Madag. Orch. 2: 26, 1941. 
Type: Comoro Islands, without proper locality. Coll. 
Humblot no. 433! (P,W). 
Kulophia tainioides Schltr. in Engl., Bot. Jahrb. 
26: 889, 1899. 
Kulophidium tainioides (Schltr.) Summerh. in 
Bull. Jard. Bot. Bruxelles 27: 408, 1957. 
Type: Mozambique, between Morumben and Mass- 
inga, region of Inhambane. Coll. Schlechter 
no. 12106! (K,Z). 
Eulophia dissimilis Dyer in FI. Pl. Afr. 27: t. 
1066, 1949. 
Eulophidium dissimile Dyer in Fl. Pl. Afr. 27: 
t. 1066, 1949, nom. altern. in obs. 


[ 264 ] 


Type: Mozambique, Lourenco Marques District, Le- 
bombo Mountains. Coll. Daintree s.n. (PRE). 
Distribution: Mozambique, Comoro Islands. 


We cannot find enough distinction between F. lonchophylla and 
E. tainioides to maintain them separately. 

Humblot no, 433 is a mixture, containing material also referable 
to O. cordylinophylla and to O. Perrieri. In the Reichenbach Her- 
barium no. 6531 the original description and drawings by Reichen- 
bach are mounted with a specimen with long, lorate leaves. This 
specimen does not belong to O. lonchophylla. We have chosen Her- 
barium Reichenbach no. 5902 as the holotype of Humblot no. 433, 
O. lonchophylla, for it agrees with the specimens under that name in 
Paris. 


Oeceoclades Lubbersiana (De Wildem. & Laurent) 
Garay & Taylor, comb. nov. 
Basionym: Hulophia Lubbersiana De Wildem. & 
Laurent in Rev. Hort. Belg. 26: 4, 1900. 
Syn.: HMulophidium Lubbersianum (De Wildem. & 
Laurent) Summerh. in Bull. Jard. Bot. Brux- 
elles 27: 397, 1957. 
Type: Zaire, Sankur. Coll. Laurent s.n./ (BR). 
Distribution: Zaire, Uganda. 
Vegetatively the plants of this species are very similar to O. atro- 
virens. The shape of the lip which is wider than long, however, is 


unique in the relationship to which also O. (latifolia and O. pandurata 
belong. 


Oeceoclades Mackenii (Rolfe ex Hemsl.) Garay & 
Taylor, comb. nov. 
Basionym: Hulophia Macken Rolfe ex Hemsl. in 
Gard. Chron. ser. 8, 12: 5838, 1892. 
Syn.: HMulophidium Macken (Rolfe ex Hemsl.) 
Schltr. in Ann. Mus. Col. Marseille ser. 3, 1: 
188, 19138. 
Type: Natal, near Verulam. Coll. McKen no. 11! (K). 
Distribution: Natal, Mozambique, Rhodesia. 


Superficially the plants of this species are rather similar to O. macu- 
lata. However, the lip which is shorter than wide and the branched in- 
florescence afford easy recognition in both the field and the herbarium, 


[ 265 ] 


Oeceoclades maculata (Lind].) Lind]., Gen. and Sp. 
Orch. Pl. 237, 18388. 
Basionym: Angraecum maculatum Lindl., Collect. 


Syn.: 


Type: 


Bot. t. 15, May 1821. 
Limodorum maculatum WLodd., Bot. Cab. 5: t. 
496, June 1821. 
Aerobion maculatum (Lindl.) Spreng., Syst. 
Veg. 3: 718, 1826. 
Eulophia maculata (Lindl.) Rehb.f. in Walp. 
Ann. 6: 647, 1868. 
Eulophidium maculatum (Lindl.) Pfitz., Entw. 
Nat. Anordn. Orch. 88, 1887. 
Graphorchis maculata (Lind].) O. Ktze., Rev. 
Gen. Pl. 2: 662, 1891. 
Brazil, without proper locality. Introduced and 
cultivated by Loddiges no. 84.10.16! (BM). 
Geodorum pictum Link & Otto, Ic. Pl. Sel. pt. 
3: 85, t. 14, July 1821. 


Lectotype: Brazil, without precise locality. Received 


Type: 


Type: 


from British Gardens and cultivated in 
Berlin. (Probably part of the original in- 
troduction by Loddiges). Holotype was 
destroyed during World War II. The 
published plate is designated here as the 
Lectotype. 

Eulophia Ledienit Stein ex N.E. Br. in Kew 

Bull. 90, 1899. 

Eulophidium Ledienit (Stein ex N.E. Br.) De 

Wildem. in Ann. Mus. Congo ser. 5, 1: 115, 

1904. 

Zaire, without precise locality. Coll. Ledien 

s.n. (WRSL). 

Julophidium Warneckeanum Kral. in Engl., 

Bot. Jahrb. 33: 70, 1902. 

West Africa, Togo, near Lome. Coll. Warnecke 

no. 196! (KX). 


[ 266 ] 


Eulophidium nyassanum Schltr. in Engl., Bot. 
Jahrb. 53: 598, 1915. 
Lectotype: Tanzania, near Mbaka Kilambo. Coll. 
Stolz no. 1909! (BM,K). 

Distribution: U.S. A.—Florida, Venezuela, Colombia, 
Guyana, Peru, Bolivia, Argentina, Paraguay, Brazil, 
Trinidad, Bahamas, Dominican Republic, W.1., Sene- 
gal, Guinee Bissau, Sierra Leone, Liberia, Ghana, Togo, 
Nigeria, Sio ‘Tomé, Gabon, Zaire, Congo- Brazzaville, 
Burundi, Sudan, Uganda, Tanganyika, Zanzibar, Pemba, 
Zambia, Rhodesia, Angola. 


The plants of all three species, O. maculata, O. monophylla, and O. 
Mackenii are very similar in appearance. Yet the proportions of the 
lip in all are sufficiently distinct to allow easy recognition. The lip 
of O. monophylla always has a distinct elongate isthmus. 


Oeceoclades maculata var. pterocarpa (Hauman) 

Garay & Taylor, comb. nov. 

Basionym: Hulophidium maculatum var. pterocarpum 

Hauman in Anal. Mus. Hist. Nat. B. 
Aires 29: 881, 1917. 

Type: Argentina, Formosa, Pilaya. Coll. Kermes 
no. 507. (BA). 

Syn.: Mpidendrum connivens Vell., Fl. Flumin. 9: t. 
44, 1851. 

Type: Brazil, Santa Cruz. Coll. Vellozos.n. No speci- 
men is known to exist. Vellozo’s original draw- 
ing in Flora Fluminensis is regarded here as 
representing the holotype. 

Distribution: Argentina, Paraguay, Brazil. 

This variety differs from the typical form in having winged cap- 

sules. It is possible that when more material is at hand, especially 
fresh flowers, this variety may prove torepresent a species sufficiently 


distinct from O. maculata. There is also a certain difference in the 
outline of the lip. 


Oeceoclades monophylla (A. Rich.) Garay & Tay- 


lor, comb. nov. 


[ 267 | 


Basionym: Angraecum monophyllum A. Rich, in 
Mem. Soc. Hist. Nat. Paris 4: 58, t. 9, 
1828. 
Syn. : Hulophidium monophyllum (A. Rich.) Schltr. in 
Ann. Mus. Col. Marseille, ser. 3, 1: 183, 1913. 
Type: Mauritius Island, without proper locality. Coll. 
Commerson s.n./ (P). 

Distribution: Mascarene Islands. 

The distinction between the plants of this species and those of O. 
maculata have already been stated above. If the distance of articula- 
tion of leaves and pseudobulb is of specific importance, as Summer- 
hayes seems to have thought, then Kulophia Ledientt now included in 
O. maculata, will probably be recognized on its own as a close relative 


of O. monophylla. 


Oeceoclades pandurata (Rolfe) Garay & Taylor, 
comb, nov. 

Basionym: Mulophia pandurata Rolfe in Journ. Linn. 

Soc. London 29: 52, 1891. 

Syn. : Lissochilus panduratus (Rolfe) H. Perr.in Hum- 
bert, Fl. Madag. Orch. 2: 29, 1841. 
Kulophidium panduratum (Rolfe) Summerh., in 
Bull. Jard. Bot. Bruxelles 27: 899, 1957. 

Type: Madagascar, near Fort Dauphin. Coll. Elhot 

no. 2546! (Ix). 
Distribution: Madagascar, Rhodesia. 


The free, truncate lateral lobes of the lip are very characteristic 
for this species. The lateral veins on the dise are papillose-ciliolate. 


Oeceoclades Perrieri (Schltr. ) Garay & Taylor, comb. 
nov. 

Basionym: MMulophidium Perriert Schltr. in Fedde 
Rep. Beih. 33: 256, 1925, not Mulo- 
phidium ambongense Schltr. 

Syn.: Mulophia ambongensis Schltr. in Ann. Col. 
Mus. Marseille ser. 8, 1: 169, t. 18, 19138. 
Lissochilus ambongensis (Schltr.) H. Perr. in 
Humbert, Fl. Madag. Orch. 2: 19, 1941. 


[ 268 ] 


Type: Madagascar, Manongarivo (Ambongo). Coll. 
Perrier no. 1654! (P). 

Distribution: Madagascar, Mozambique. 

The long-acuminate, lorate leaves without a distinct petiole dis- 


tinguishes the plants of this species from the related O. quadriloba, 
O. sclerophylla and O. analavelensis. 


Oeceoclades petiolata (Schltr.) Garay & Taylor, 
comb. nov. 
Basionym: Mulophia petiolata Schltr. in Ann. Mus. 
Col. Marseille ser. 3, 1: 175, t. 18, 1913. 
Syn.: Mulophidium petiolatum (Schltr.) Schltr. in 
Fedde Rep. Beih. 33: 256, 1925. 
Lissochilus petiolatus (Schltr.) H. Perr. in 
Humbert, Fl. Madag. Orch. 2: 25, 1941. 
Type: Madagascar, Manongarivo (Ambongo). Coll. 
Perrier no. 478 bis! (P). 
Distribution: Madagascar. 


Vegetatively the plants of this species are practically identical with 
those of O. alismatophylia, but the shape of the lip is very different 
proportionately. From the related O. lonchephylila it differs in having 
a pandurate lip with three thickened veins in front of the calli. 


Oeceoclades quadriloba (Schltr.) Garay & Taylor, 
comb, nov. 
Basionym: MWulophia quadriloba Schltr. in Ann. Mus. 
Col. Marseille ser. 8, 1: 176, t. 12, 1918. 
Syn.: Hulophidium quadrilobum (Schltr.) Schltr. in 
Fedde Rep. Beih. 33: 256, 1925. 
Lissochilus quadrilobus (Schlitr.) H. Perr. in 
Humbert, Fl. Madag. Orch. 2: 30, 1941. 
Type: Madagascar, Manongarivo (Ambongo). Coll. 
Perrier no. 1696! (P). 
Distribution: Madagascar, Rhodesia. 


The unique structure of the lip in these plants is reminiscent of 
those belonging to the Asiatic genus Grosourdya. The vesicular, hang- 
ing spur is longer than the reflexed midlobe of the lip. 


[ 269 | 


Oeceoclades Rauhii (Sengh.) Garay & Taylor, comb. 
nov. 
Basionym: Mulophidium Rauhiu Sengh. in Die Orch- 
idee 24: 61, 1973. 
Type: Madagascar, south of Anivorano. Coll. Rauh 
& Senghas no. 22865 (HEID). 
Distribution: Madagascar. 
Closely related to O. boinensis from which it differs in having liner- 
lanceolate sepals and petals and an equally four-lobed lip. 


Oeceoclades roseovariegata (Sengh.) Garay & Tay- 
lor, comb. nov. 
Basionym: Mulophidium roseovariegatum Sengh. in 
Adansonia ser. 2, 6: 561, 1967. 

Type: Madagascar, near Diégo-Suarez, ‘‘Montagne 
des Francais’’. Coll. Rauh & Buchloch no. 
7985 (HELD). 

Distribution: Madagascar. 


As it was stated above, this species is closely allied to O. gracillima. 
As a matter of fact, the plants were already known to Schlechter 
through a collection by Perrier no. 16224! (P), and was regarded by 
him as an undescribed species. The Perrier specimen has Schlechter’s 
original drawings of the floral parts attached to the sheet. It was also 
collected on ‘‘Montagne des Francais’’. Perrier identified it as Eulo- 
phidium gracillimum var., but cited it without a varietal name in his 
Orchids of Madagascar. 

The photographs of O. roseovariegata in Die Orchidee 18: 24, 1967 
show the spur as being distinctly bilobed. This is apparently due to 
the angle in photographing, revealing the ventrally compressed and 
slightly grooved tip. 


Oeceoclades Saundersiana (Rchb.f.) Garay & Tay- 
lor, comb. nov. 
Basionym: Hulophia Saundersiana Rehb.f. in Bot. 
Zeit. 24: 378, 1866. 
Syn.: Graphorchis Saundersiana (Rehb.f.) O. Ktze., 
Rev. Gen. Pl. 2: 662, 1891. 
Mulophidium Saundersianum (Rehb.f.) Sum- 


merh. in Bull. Jard. Bot. Bruxelles 27: 401, 
1957. 

Type: Africa, West Coast, without precise locality. 
Coll. Mann s.n./ (W). 

Lissochilus barombensis Kral. in Engl., Bot. 
Jahrb. 17: 52, 1898. 

Type: Cameroun, Barombi. Coll. Preuss no. 546! (Kx). 
Kulophia Bierleri De Wildem., Not. Pl. Util. 
Congo 1: 311, 1904. 

Type: Zaire, Coquilhatville. Coll. Bierler s.n. (BR). 
Hulophia Mildbraedi Wrz). in Engl., Bot. 
Jahrb. 43: 339, 1909. 

Type: Zaire, Ruwenzori Range, Semliki Plains. Coll. 
Mildbraed no. 275! (Kx). 

Distribution: Sierra Leone, Liberia, Ivory Coast, 

Ghana, Nigeria, Cameroun, Gabon, 
Zaire, Uganda, Kenya, Tanzania, Zam- 
bia, Angola. 


The long cylindrical pseudobulbs with two leaves and the equally 
four-lobed lip of rather large flowers easily identify the plants of this 
species. 


Oeceoclades sclerophylla (Rchb.f.) Garay & Tay- 
lor, comb. nov. 
Basionym: MWulophia sclerophylla Rchb.f. in Flora 68: 
542, 1885. 
Syn. : Mulophidium sclerophyllum (Rehb. f.) Summerh. 
in Bull. Jard. Bot. Bruxelles 27: 402, 1957. 
Type: Madagascar, Forét d’Ankaye. Coll. Humblot 
s.n.! (W). 
Hulophia Elhoti Rolfe in Journ. Linn. Soc. 
London 29: 52, 1891. 
Lissochilus Elliot (Rolfe) H. Perr. in Hum- 
bert, Fl. Madag. Orch. 2: 47, 1941. 
Type: Madagascar, near Fort Dauphin. Coll. Elliot 
no. 2424! (Kx). 


[ 271 ] 


Among the plants with long, linear leaves O. sclerophylla may be 
compared with O. analavelensis, but the two-leaved pseudobulbs and 
the differently proportioned lips readily separate the two. Vegeta- 
tively it also resembles O. quadriloba. 


Oeceoclades seychellarum (Rolfe ex Summerh.) 
Garay & Taylor, comb. nov. 
Basionym: Mulophia seychellarum Rolfe ex Summerh. 
in Bull. Mise. Inf. Kew 3638, 1928. 
Syn.: Mulophidium seychellarum (Rolfe ex Summerh. ) 
Summerh. in Bull. Jard. Bot. Bruxelles 27: 
402, 1957. 
Type: Seychelles, Mahé, Cascade Estate. Coll. 
Thomasset no. 88! (K). 


Distribution: Seychelles. 

Vegetatively the plants of O. seychellarum are identical with those 
of O. lanceata. The two may be kept apart on account of the differ- 
ences in the floral structures, especially in the shape and proportion 
of the lip as shown in the key. 


Oeceoclades spathulifera (H. Perr.) Garay & Tay- 
lor, comb. nov. 

Basionym: Hulophia spathulifera H. Perr. in Bull. 
Soc. Bot. Fr. 82: 157, 1935, as J. spatu- 
lifera, sphalm. 

Syn.: Lissochilus spathulifer (H. Perr.) H. Perr. in 
Humbert, Fl. Madag. Orch. 2: 33, 1941. 
Hulophidium spathuliferum (H. Perr.) Sum- 
merh. in Bull. Jard. Bot. Bruxelles 27: 403, 
1957. 

Type: Madagascar, Ambongo-Boina. Coll. Perrier 
no. 15930 (P). 


Distribution: Madagascar. 


The distinctly spathulate sepals and petals combined with a glo- 
bose spur easily separates the plants of this species from the related 
O. calcarata and O. Hebdingiana. 


[ 272 ] 


Oeceoclades ugandae (Rolfe) Garay & Taylor, comb. 
nov. 
Basionym: Hulophia ugandae Rolfe in Bull. Mise. 
Inf. Kew 839, 1913. 
Type: Uganda, Mabira Forest. Coll. Brown no. 443! 
(Kx). 
Distribution: Zaire, Ghana, Uganda. 


Summerhayes considered this species to be synonymous with O. 
latifolia. As a matter of fact the shape of the lip superficially looks 
very much alike in both. Yet vegetatively the plants of these two 
species are closer to other members than to one another. The lip of 
O. ugandae is also provided with a pair of gibbosities at the junction 
of the lateral and midlobes, this character is absent in O. latifolia. 


Oeceoclades zanzibarica (Summerh.) Garay & Tay- 
lor, comb. nov. 
Basionym: Mulophidium zanzibaricum Summerh. in 
Bull. Mise. Inf. Kew 417, 1927. 
Type: Zanzibar, without precise locality. Coll. Last 
s.n.! (KX). 
Distribution: Zanzibar, Pemba. 


The lanceolate, long-attenuate leaf is rather unique in the genus. 
Florally the plants of this species are closest to those of O. alismato- 
phylla from Madagascar, but differ from one another in the shape of 
the sepals and petals and in the proportions of the lip. 


[ 273 ] 


Excluded Species 
. faleata (Thunb. ) Lindl. = Neofinetia falcata (Thunb.) Hu 


. flexuosa Lindl. =Cleisostoma ramosum (Lindl. ) Hook.f. 

. funalis (Sw.) Lindl. = Dendrophylax funalis (Sw.) Benth. ex Rolfe 
. gracilis (Thou.) Lind]. =Chamaeangis gracilis (Thou. ) Schltr. 
javanica Teijsm. & Binn. = Hymenorchis javanica (T. & B) Schltr. 
Lindleyana Regel. = Neofinetia falcata (Thunb. ) Hu 

. Lindleyi Regel = Neofinetia falcata (Thunb. ) Hu 

. paniculata Lindl. = Robiquetia succisa (Lindl. ) Seidenf. & Garay 
. parviflora (Thou.) Lind], = Angraecopsis parviflora (Thou. ) Schltr. 
. pusilla Lind]. =Saccolabiopsis pusilla (Lindl.) Seidenf. & Garay 
. Retzii Lindl. =Chiloschista pusilla (Retz) Schltr. 

. tenera Lindl. = Trichoglottis tenera (Lindl.) Rchb.f. 


SOS on ee O46 5 5 


The following list of names constitutes the Section Pulchrae Krz]l. 
of the genus Eulophia (Gard. Chron. ser. 3, 22: 262, 1897). Some 
of these names have been referred to the genus Eulophidium ( = Oeceo- 
clades) previously. 

Eulophia gracilis Lindl. in Bot. Reg. 9: t. 742, 1823. 

Eulophia emarginata Bl., Fl. Java, n.s. 1: 152, 1858. 

Eulophia guamensis Ames in Philipp. Journ. Sci. Bot. 9: 12, 1814. 
Eulophia macrostachya Lind|,, Gen. and Sp. Orch. Pl. 183, 1833. 
Eulophia megistophylla Rchb.f. in Flora 68: 379, 1885. 

Eulophia minimiflora Kral. in Not. Syst. 4: 137, 1928. 

Eulophia novo-ebudae Krzl. in Bull. Soe. Bot. Fr. 76: 301, 1929. 
Eulophia pulchra (Thou.) Lind]., Gen. and Sp. Orch. Pl. 182, 1833. 
Eulophia Rourii Kral. in Sarasin & Roux, Nova Caled. 1: 82, 1914. 
Eulophia silvatica Schltr. in Engl., Bot. Jahrb. 53: 5 

Eulophia striata Rolfe in Journ. Linn. Soc. London 29: 53, 1891. 
Eulophia Wendlandiana Krzl. in Gard. Chron. ser. 3, 22: 262, 1897. 
Lissochilus ambrensis H. Perr. in Not. Syst. 14: 159, 1951. 


BOTANICAL MUSEUM LEAFLETS 


HARVARD UNIVERSITY 


CAMBRIDGE, MASSACHUSETTS, NOvEMBER 30, 1976 VoL. 24, No. 10 


MANICARIA SACCIFERA AND ITS CULTURAL 
SIGNIFICANCE AMONG THE WARAO INDIANS 
OF VENEZUELA 


BY 
JOHANNES WILBERT 


Within the tropical and subtropical belt that circles the 
earth, palms can truly be counted among the best friends of 
man. In several regions, the various parts of the palm are so 
thoroughly exploited for purposes of food, drink, basic ma- 
terials, tools, and utensils that it assumes a pivotal position 
in the cultural life of the people. In South America outstand- 
ing examples are Astrocaryum, Euterpe, Guilielma, Jessenia, 
Mauritia, Maximiliana, Oenocarpus, Orbignya, and Syagrus. 
In addition, the Indians are known to utilize the products of 
at least a dozen other genera of palms.’ 

This paper focuses on Manicaria saccifera, the temiche 
palm, and its cultural importance among the Warao Indians 
of Venezuela (PLATE LXVI). Ethnobotanical information 
on this genus, except for descriptions in floras, is slight, and 
the fact that Manicaria saccifera is used as a source of sago 
has, heretofore, gone unrecorded in the scientific literature.” 

As an ancient food-quest activity of man, the recovery of 
palm starch “appears to be a pantropical phenomenon that is 
most highly developed on the mainland of Southeast Asia and 
in the West Pacific. In that region starch extraction involves, 
principally, palms of the genus M etroxylon, Sago Palms 
(Burkill 1935: 1460-1462; and Barrau 1959: 151-159). Palms 


[ 275 ] 


PLATE LXVI 


Manicaria saccifera in the forest of the Intermediate zone of the 
Orinoco Delta. 


[ 276 ] 


of other genera are also used to a lesser extent” (Heinen and 
Ruddle 1974: 116). : 

In South America, the Indians recover starch from at least 
four genera of palms: Syagrus, Copernicia, Mauritia, and 
Manicaria, the most important being Syagrus and Mauri- 
tia.” Syagrus Romanzoffianum and Copernicia cerifera are 
exploited for sago by the Tupi-speaking Guayaki Indians of 
Paraguay and by several Guaranian and non-Guaranian 
tribes of the Gran Chaco. Mauritia flexuosa and Manicaria 
saccifera are utilized for sago recovery by the Warao Indians 
of the Orinoco Delta in Venezuela and in the adjacent east- 
erly regions of Guyana. 

Syagrus Romanzoffianum, the Paraguayan coconut or 
queen palm, has a southeasterly distribution in South Amer- 
ica, where it is known by its Guarani name pindo. The Gua- 
yaki call the palm toi (tdi, tdi) and refer to its sago as kraku 
(Cadogan 1960). Because of the many uses to which its 
different parts are put, Syagrus Romanzoffianum plays an 
outstanding role in the lives of the autochthonous and rural 
populations of this part of the New World. In fact, for the 
different Indian tribes of the area it represents probably the 
most important economic plant of their environment. 

The extraction of sago from Syagrus Romanzoffianum 
and Copernicia cerifera is known to be practiced in South 
America by such tribes as the Guayaki, Mbya (Caingua), 
Kaingan, Toba, Lengua, and Chamacoco. But it is highly 
probable that this practice was diffused much more widely 
in earlier, pre-agricultural times. Writes Vellard (1934-35: 
240-241) concerning the Guayakti: 

. it is the flour extracted from the pindo palm (Cocos Roman- 
zoffiana) which, along with the game and honey, constitutes their 
basic diet. . . . To prepare the flour . . . the Guayaki cut open the 
trunk with an axe. The fibers are pounded and crushed on the 
spot with the help of an old piece of bow or with the back of 
the axe, then they are roughly strained on a square frame with 
loose straw or lamellas of bamboo bark. After being moistened 


with water the flour is used for making cakes which are eaten raw 
or dried near the fire. 


[ 277 ] 


Vellard (ibid., p. 240) points out that krakw starch is pre- 
pared not only by the Guayaki but also by the Mbya, a sub- 
group of the Guarani-speaking Caingua. They fall back on it 
as an emergency food. “To obtain the starchy pith of palm 
trees, the Mbya extracted the long fibers imbedded in starch 
from the lower part of the trunk. They either pounded them 
in a mortar and sucked them or else dried them on a platform 
in the sun or over the fire, pounded them, sifted them through 
a net, and then made them into loaves or cakes” (Métraux 
1946a: 262). Even the Paraguayans resorted to eating kraki 
“after the disastrous war against the Triple-Alliance (1866- 
1870)” (Vellard ibid. p. 240). 

For the Kaingin (Caingang), a non-Guarani-speaking 
tribe of southern Brazil, the sago of Syagrus Romanzoffi- 
anum was an important food before it was replaced by man- 
ioc flour. (The Indians crushed the pith in a mortar and 
sifted the flour before roasting it in a pan, just as is done now 
with manioc flour.) (Métraux 1946b: 445-453. ) 

Métraux (1946a: 248, 261) also describes the Toba, the 
Lengua, and the Chamacoco tribes of the Gran Chaco as 
recovering the palm starch of Copernicia cerifera. The Toba 
pound the pith in a mortar and boil it as a mush, whereas the 
Lengua grate it into flour for cakes. Carandaipe starch is a 
principal vegetable food for the Chamacoco,. 

The best documented case by far of palm-starch extraction 
for any South American Indian tribe comes from the region 
of the Warao, where chroniclers, missionaries, travellers, and 
anthropologists have become aware of its existence and 
where, on numerous occasions over the past twenty years of 
intermittent field work, I witnessed at first hand the process 
of recovery of sago from Mauritia.”. The ethnobotanical data 
available on this palm are too abundant to be treated here. 
Suffice it to say that Mauritia sago, ohidu aru, has been the 
staple food for most Warao until very recently, when it was 
supplanted by ocumo (Xanthosoma sagittifolium) and, to a 
lesser extent, by manioc (Manihot esculenta). 

The practice of extracting sago from Manicaria saccifera 


[ 278 ] 


(yahuhi aru) came to my attention only recently, and I 
was able to witness the procedure for the first time in the 
summer of 1975. 

In a process that may have taken more than a thousand 
years, the Warao have adapted their life and culture to the 
rather difficult outer world of the Orinoco Delta, situated in 
eastern Venezuela between 8° and 10° north latitude and 
between 59° and 62° west longitude. The Delta is a large 
fan of alluvial deposits occupying 17,000 square kilometers 
and bounded on the south by the Orinoco River proper and 
on the west by the Manamo which branches off the main 
river at Barrancas, where the apex of the Delta is situated. 
Most of this low-lying triangle is a tidal swamp lacking in 
dry ground and stone and extending inland from 50 to 100 
kilometers (Liddle 1928: 20-24). 

Manicaria does not grow throughout this vast area but is 
restricted mainly to regions immediately within the Inter- 
mediate Delta zone behind the Lower Delta, most of which 
is covered by a belt of mangrove. Typical of such environ- 
ments, the soil is almost always inundated, thus providing 
along the perimeter of the swamp the ideal condition for the 
pioneering Rhizophora Mangle. The feature characteristic 
of mangrove forests of growing on the periphery while dying 
at the core is of primary importance to the Warao. The clear- 
ing that results from the decomposing inner parts of the 
mangrove forest is invaded by many kinds of trees important 
to the Indians, among them useful palms such as the Mau- 
ritia, Euterpe, and Manicaria. Thus, within and behind the 
coastal mangrove belt of the Lower and Intermediate Delta 
zones there developed the echonich of a palmetum that has 
amply served the Warao as an abundant food basket and as a 
secure home, 

The average mean temperature of the Delta is 26° C, the 
humidity 60 to 80 percent. The rainy season lasts roughly 
from May to October and the dry season from November to 
April, but rainy days with more or less intensive showers oc- 
cur throughout the year. The annual precipitation ranges 


[ 279 ] 


from 100 to 200 centimeters (Heinen and Lavandero 1973: 
4-11), 

Twice daily the tide washes over the palmetum, encourag- 
ing the growth of the trees. The annual flooding of the Ori- 
noco is felt only indirectly in the Intermediate Delta; the 
flooding of the palm groves during the wet season is actually 
due to rainfall. In the dry season, when the waters of the 
Orinoco recede, sea water penetrates into the Intermediate 
Delta, salinates the rivers, and temporarily causes a potable 
water problem. Within the palmetum, Manicaria appears to 
seek out not only places under the influence of the tides but 
also those exposed to the northeasterly trade winds that 
sweep over the Delta almost incessantly. 

Warao culture is particularly adapted to life in this palm- 
rich environment. For the pre-agricultural Warao, the sago 
primarily of Mauritia and secondarily of Manicaria provided 
the staple food as well as a superabundance of edible fruits 
during much of the year. The Euterpe (E. edulis), too 
yielded fruit and an especially tasty and rich palmito. Mani- 
caria milk and Mauritia (unfermented) wine helped solve 
any drinking water problem, and the fat grubs of the palm 
borer (Ryncophorus palmetum), collected in overwhelming 
quantities from fallen Mauritia and Manicaria, was an impor- 
tant added source of protein to their diet. That the palms 
are a blessing is clearly recognized by the Warao, and these 
plants permeate their entire culture: its technological, socio- 
economic, and religious systems. The Indians refer to Mau- 
ritia sago, especially in combination with fish, as the “true 
food” of man; Manicaria, to a lesser degree, falls into the 
same category. Sago was more than a vital source of human 
sustenance; it came to be elevated to a position of ritual 
significance which has helped the Warao to cope psycho- 
logically with the hardships imposed upon them by a refuge 
environment little amenable to human life and culture. 

Other palms found in the Orinoco Delta, but mostly out- 
side the palmetum of the Lower and Intermediate zones, are 
Astrocaryum, Jessenia, Maximiliana, Socratea and others. 


[ 280 ] 


The study of Manicaria began in 1791, when the German 
botanist Joseph Gaertner (1791: 468, Pl. 2) founded the 
genus on the basis of the spathe, spadix, rachilla, and flowers 
of the plant (PLare LXVII). Apparently, its trunk, foliage, 
and fruit remained largely unknown to the scientific world 
until sixty-two years later, when Wallace (1853: Plates II 
and XXVI) furnished his illustration of this “unique and 
handsome palm” (PLate LXVIII). Wallace and other field 
workers have added much information to the original de- 
scription,’ although the genus, its speciation and distribution 
remain relatively little understood even today.’ 

Standley and Steyermark (1958: 271) have published a 
concise summary of our present knowledge on the genus: 


MANIcARIA Gaertner 


Reference: Burret, Notizbl. Bot. Gart. Berlin 11: 389. 1928. 


Plants very robust, tall, or low, unarmed, the caudex stout, 
annulate, often curved or flexuous, covered with old leaf sheaths; 
leaves terminal, very large, suberect, lanceolate, acute, plicate- 
nerved, serrate at first and finally pinnatisect, the costa thick and 
stout, the petiole slender, the sheath cleft, its margins with many 
coarse fibers; spadices several, erect-spreading, tomentose, the 
branches strict, rather thick, foveolate; spathes 2, the upper fusi- 
form, terete, mucronate, fibrous, tardily rupturing; bracts subulate; 
flowers monoecious, borne in the same spadix, this inserted among 
the leaves, simply branched, the flowers immersed in pits in the 
branches, the upper ones staminate, crowded, the lower ones 
scattered, pistillate; staminate flowers oblong trigonous, the sepals 
ovate-rounded, coriaceous, with scarious margins, imbricate, the 
petals thick-coriaceous, obovate-oblong, valvate; stamens 24-30, 
the filaments filiform, connate at the base, the anthers narrowly 
linear, erect, bifid at the base, emarginate; pistillate flowers larger, 
ovoid, the perianth little enlarged after anthesis, the sepals 
rounded, their margins finally lacerate, broadly imbricate, the 
petals longer, covolute-imbricate at the base, acute and valvate 
at the apex; ovary sulcate, 3-celled, the stigmas 3, sessile; fruit 
large, globose, 1-seeded, or depressed-globose and 2-3-seeded, 
the stigmas terminal, the pericarp corticate, the cortex corky, an- 
gulate-echinate, the endocarp vitreous-crustaceous, fibrous within; 


[ 281 ] 


PLATE LXVII 


Spathe, spadix, rachilla, and flowers that served Gaertner to estab- 
lish the genus Manicaria, 1791. 


[ 282 ] 


PLATE LXVIII 


Teod A Vest ng 


Waa 


MANICARIA SACCIFERA Ht 40F 


1853. 


> 


First depiction by Wallace of Manicaria 


[ 283 ] 


seeds globose, erect, the hilum oblou.g, the testa very hard, the 
branches of the raphe closely reticulate and involving the seed, 
the endosperm corneous, uniform, the embryo basal. 


The genus has become known from Central America 
(Bailey 1943: 392-393; Standley and Steyermark 1958: 271— 
273), Trinidad (Bailey 1933: 409-413) and northern South 
America: from Colombia, the Orinoco Delta, the Guianas 
( Wessels Boer 1965a, b) to the mouth of the Amazon ( Burret 
1928: 389). It also occurs on the Rio Negro and the Upper 
Amazon ( Wallace 1853: 70). 

A very complete description of the species Manicaria 
saccifera was written by Wessels Boer (1965a: 21) PLATE 
LXIX: 


Trunk solitary, up to 6 m. tall, about 3 dm. in diameter, in the 
upper part covered with dead leaf-bases, at base with prominent 
leaf-scars. About 10 contemporaneous suberect leaves; dead 
leaves persistent for some time and hanging down on the trunk; 
sheath with fibrous ventral part enclosing young leaves about 7 
dm. long; petiole stout, about 12 dm. long and 8 cm. in diameter, 
grooved, leaf-blades very large, simple or irregularly pinnatisect 
through the action of the wind, also in leaves of juvenile plants, 
up to 72 m. long and 23 dm. wide, bifid at apex, margin serrate; 
about 120 primary veins, 3-4 cm. distant at the middle of the 
blade, secondary veins inconspicuous; petiole, costa, and the 
lower surface of the blade at first more or less brown-tomentose, 
soon glabrescent. 

Spadices almost erect, about 17 dm. long, with 2 spathes; outer 
spathe about 7 dm. long, flattened, invisible between leaf-sheaths, 
inner spathe about 11 dm. long, fusiform, mucronate, consisting 
of densely interwoven fibers without any suture, enclosing the in- 
florescence completely till long after anthesis; peduncle about 10 
dm. long, rachis about 6 dm. long with up to 45 simple rachillas 
or rarely a few rachillas bifurcate, several large bracts along the 
peduncle within the inner spathe, smaller bracts at the base of 
the rachillas. Male flowers densely crowded in the upper part of 
the rachillas (and 2 laterally adjacent to each female flower), 
sunken in small pits and subtended by bracts 7-12 mm. long; 
sepals ovate, imbricate, 3-4 mm. long, petals lanceolate, valvate, 
ligneous-incrassate, 6-7 mm. long; stamens many (20-34), 
densely congested, filaments about 1% mm. long, anthers 3 mm. 
long, the central ones usually misshapen. Female flowers few, 


[ 284 ] 


PLATE LXIX 


Depiction in Standley and Steyermark (1958: Fig. 45) of details of 
leaf, flower, and spathe of Manicaria saccifera. 
(Courtesy Cuicaco Natura History Museum) 


[ 285 ] 


near the base of the rachillas between 2 male flowers, subtended 
by bracts; sepals ovate, imbricate, 7 mm. long, 8 mm. wide, petals 
ovate, valvate, acute at apex, ligneous-incrassate, 10 mm. long, 
6 mm. wide; pistil globose, 4 mm. in diameter, stigma 3-fid, sessile, 
erect, 3-4 mm. long, strongly papillose. Fruit depressed-globose, 
3-seeded or by abortion 1- or 2-seeded; exocarp corky, angulate- 
echinate, tubercles rather hard and not easily rubbed off; seed 
globose, about 4 cm, in diameter; embryo basal . . . 


In this account of the genus, the most recent, Wessels Boer 
goes on to identify four species of Manicaria, very con- 
spicuous and abundant swamp plants. The Manicaria occur 
in forests interspersed with other trees but also form colonies, 
or temichales, of great density. Manicaria saccifera is known 
in Venezuela as temiche (timiche, timiti), an Arawakan 
noun; in Guyana as truli, a Cariban term; and in Brazil as 
ubussti, a Tupian word meaning “big leaves” (Civrieux 1957: 
195-232). The Warao refer to it poetically as yahuhi 
(yawihi) meaning “plumes of the sun,” descriptive of the 
leaves that look like giant bird feathers. 


The Origin Myth of the Temiche Palm 


Long ago there was an old woman who followed her 
husband to live in the lowlands of the Delta. Life was easier 
for the old couple there than where they had come from, 
and they greatly enjoyed the cool water of the bogland and 
the sea breeze that incessantly fanned their new home. The 
name of the old woman was Yahuhi. 

As time went by, the woman felt a strange transformation 
taking place all over her body. First her eyes began to clear 
up so that she could see well again. Then the wrinkles in 
her face disappeared, her body firmed up like that of a girl, 
and her complexion became healthy and youthful looking. 
Even the voice of the old woman changed back to that of a 
maiden, and she began to sing with happiness over her re- 
gained youth. 


The husband of the woman was equally taken by surprise 


[ 286 ] 


and wondered what the cause of this miraculous transforma- 
tion might have been. 

“I have changed because my body was exposed to the 
cool northwind,” said the woman. “Let’s remain here for- 
ever.” 

One day, the man told his youthful wife that he had to 
leave the house for a short while. He wanted to go to the 
field and do some gardening. 

“Fine,” said the woman. “I shall cook dinner and wait for 
you. But don't be later than you said.” 

Time passed, and the man failed to return within the set 
period of time. While his wife was waiting, she suddenly 
saw a handsome young man approaching the house from the 
North. He greeted her kindly and wanted to know where 
her husband had gone. “He has gone to the field and is 
overdue.” 

So then the visitor took advantage of the man’s absence 
and seduced the youthful woman. 

When the husband finally got home, the suitor had long 
since left the house, but the husband noticed that something 
was wrong. Questioning his wife, she finally confessed that 
a young man had visited and embraced her during his 
absence. It was his fault, she said. He had left her alone 
for so long. 

This made the old man very angry. He prepared a rope 
and whipped the poor woman so mercilessly that marks 
began covering her body from head to foot. Days went by, 
and the woman became very ill. Because of the pain, she 
could neither sleep nor eat and, finally, she died. 

Since there were no people living in the neighborhood who 
might have helped the old man bury his wife, the husband 
tied the corpse to a pole in an upright position. It looked as 
if the woman was just standing there, alive. And even after 
a full moon had passed, the woman’s body still looked youth- 
ful and uncorrupted. 

After that, however, the dead woman began to change into 
a tree. The husband left her, and upon looking back a final 


[ 287 ] 


time, saw that his wife had become a temiche palm. He said 
to himself: “Once the Warao will come to be on this earth, 
they will have to call this palm yawuhi, because that was 
her name as a woman.” 

The etiological intent of this simple narrative is clear: a 
prolific palm bearing fruit practic sally continuously is iden- 
tified with a fertile young woman. Her fertility is mirac- 
ulously caused by the iad of the north, whence also her 
youthful lover puts in an appearance. 

For a Warao listener, the introduction of the northwind 
heralds doom. While the cool nortes in the Delta are certainly 
invigorating winds, they also bring catarrh and other res- 
piratory ailments to the Warao. Furthermore, they blow 
from the direction where Haburi, the culture hero, lives in 
a world mountain-tree. As a youth, he had unwittingly 
seduced his own mother. The association of Haburi with 
the woman’s paramour from the north is most certainly not 
lost on a Warao listener. In addition, Haburi himself had 
been made miraculously youthful and mature through the 
agent of an old frog-woman, who wanted him for a lover. 
Both the youthful temiche woman and Haburi were placed 
into a fateful triangle by virtue of their newly acquired 
sexual prowess. Adultery and incest were the inevitable 
consequences, since the partners of both the temiche-maiden 
and Haburi were infertile and no match. Consequently, the 
woman transformed into a palm as did Haburi into a world 
tree, 

This sacrifice of metamorphosis results in enormous bene- 
fits for mankind: from the transformed woman, the ever 
producing temiche palm; and from the transformed hero, 
the dugout canoe, a sine qua non of human life in the Delta. 
So while the Warao listener can predict the tragedy that will 
inevitably result from the action of the dramatis personae, 
he can also anticipate the great benefits that will accrue to 
him from this primordial drama of the temiche-maiden, the 
old husband, and her paramour from the North. 

The myth also explains why the temiche favors coastal 


[ 288 ] 


swamps with their tidewaters and sea breeze. Besides ex- 
plaining the remarkable fertility of the palm, subject not to 
an annual flowering and fruiting season but to continuous 
yield and the swamp habitat of the temiche, the myth also 
offers an explanation for the exceptionally prominent leaf 
scars (yd esoara) that cover the entire stem of the palm. 
Similar explanations are given in Warao mythology for the 
rings of Euterpe and of trees like Calophyllum. Finally, as 
we shall see, different parts of the palm are used as med cue 
against respiratory illnesses. This blissful property of the 
plant may possibly find its explanation in the love of the 
tree-maiden for the nortes that commonly cause such ills. 


Utilization of the Palm 


The leaves. It is well to commence the discussion of the 
cultural significance of Manicaria for the Warao Indians with 
the plant’s most outstanding characteristic, its leaves. The 
“plumes of the sun,” yahuhi, as the Warao call them, are 
the largest entire leaf among palms and the lar gest in the 
plant kingdom. The palm seudied for purposes of this paper 
had seventeen contemporaneous sub-erect leaves with two 
persistent dead leaves hanging down. The informant hap- 
pened to know that the palm was between 30 and 35 years 
old. Along their entire axis, large leaves of the Manicaria 
saccifera measured 5-8 m. long ae 1.5-1.8 m. wide, with 
petioles measuring 1.20-2 m. Spiaaa (1968: 111) reports 
leaves 9 m. in length, and leaves of 10 m. are frequently 
mentioned in the literature. Through exposure to wind, the 
blades of large outer leaves eal to become irregularly 
pinnatisect, but younger inner blades remain undivided. 

It is precisely this latter quality coupled with their size 
that make Manicaria leaves especially suited for house thatch 
wherever the plant grows (PLare LXX). As Im Thurn 
(1967 [1883]: 209) observed, “each gigantic undivided leaf 
of the troolie palm ( Manicaria saccifera) is really a shelter in 
itself; and a few of these laid, without further preparation, 


[ 289 ] 


PLATE LXX 


Warao houses with Manicaria thatch. Mauritia flexuosa in the 
background. (Courtesy P. T. Furst) 


[ 290 ] 


so as to overlap like tiles, make a most perfect roof. Indeed, 
before corrugated zinc was introduced for the purpose, a 
large trade was carried on between the Indians and the 
planters on the coast in these troolie-leaves, with which most 
of the buildings on the sugar estates were thatched.” 

For weather protection, the Warao implant single leaves 
or a whole line of them into the soft ground near their work- 
ing areas. They also cover their heads with leaf segments 
when traveling on foot or by boat, calling these makeshift 
umbrellas aroko a kuasimara, leaf capes. 

To thatch their houses, the Indians fold the leaves in half 
along the rachis and lash them in overlapping fashion, each 
rib 25 cm. from the next, vertically onto the infra-structure 
of the roof, “so that each frond forms a long tile reaching 
from ridge to eaves” (Spruce 1908, 1: 59) (PLarE LXXI1). 
The house of my main informant had been covered in this 
way in 1969 and did not begin to leak until 1975, testimony 
of the durability of temiche thatch. To keep the rain from 
drifting in with the sea breeze, the Warao install a screen of 
temiche (dara yawihi) on the windward side of their other- 
wise wall-less houses * (PLare LXXII). Sometimes they 
construct a tunnel-like roof over the midsection of their dug- 
out canoes to protect themselves from the weather during 
long journeys. 

An ingenious naval invention is making sails (yawihi 
wera) from large Manicaria leaves. On the open windswept 
canos of the Delta, Warao canoes go by at high speed under 
full sail. Two or three crew members each hold up a 
Manicaria leaf for sails, bracing it at the bottom against the 
foot and holding it with one arm (PLATE LXXIII). A helms- 
man keeps the course by means of a paddle held vertically 
as a rudder. I have clocked canoes 6 m. long with two 
paddlers but no sails going 3 kph, their full speed. Canoes 
with temiche “sails” go that fast, or faster, and, of course, 
for a longer period of time. Consequently, in terms of 
primitive navigation, the yawihi wera of the Warao repre- 
sents critical navigational tackle. Despite its Spanish-derived 


[ 291 | 


[ 26a | 


Detail of roofing of a Warao house. (Courtesy P. T. Furst) 


IXX'] 4LVId 


PLATE LXXII 


Detail of weather screen on the windward side of a Warao house. 
(Courtesy P. T. Fursr) 


[ 293 ] 


| £62 | 


Three canoes with some fourteen passengers and a heavy load of fire wood are being propelled by 


holding up three leaves of Manicaria saccifera. Warao Indians of the Intermediate zone ‘of the Orinoco 
— Ce Oe ee > i pe en | 


HIXX'] 2LV 1d 


name (wera=vela, sail), I wonder whether, in view of its 
simplicity, sailing by yawihi wera does not antedate the 
arrival of the white man in northeastern South America. 
Since Manicaria is ubiquitous in the Lower and Intermediate 
Delta zones, a “sail” can be picked up at practically any point 
of departure and simply tossed away upon arrival at the 
destination. In other words, Manicaria leaf-sails are handy, 
free, and uncomplicated. 

Manicaria leaves, or certain parts, are put to other uses 
by the Warao Indians. The tips of the leaves are improvised 
for use as fans (yami); several, 2 m. long pieces of rachises 
of leaves are tied together in the form of a Venetian blind to 
serve as fish weirs (noba); sections of midrib are rubbed 
together to produce fire by rotation (Im Thurn 1967: 257). 
The “plumes of the sun” represent a materia prima of great 
importance to Warao technology. 

A final comment on the etymology of the Warao term 
“plumes of the sun.” It is derived from far more than the 
shape of leaves that flicker in the sun (as some authors seem 
to suggest). The Manicaria leaves obviously resemble over- 
sized bird feathers; but why they are linked to the sun is 
less evident. Manicaria, like many other palms, are closely 
connected with a symbolism of light and darkness, day and 
night, and I discuss this aspect below. 

The spathe. Next in importance are the pouch-like spathes 
that cover the entire inflorescence and the large pendant 
infructescence (PLATE LXXIV). The brown spathes from 
which the genus and the species derive their names ° are from 
40 to 60 cm. long, “of fine, closely woven texture, and are 
used by the natives to make soft brown caps without seams 
or joinings” (McCurrah 1960: 129). The Warao refer to 
these hood-like caps as yasi nona. After carefully peeling 
the spathe off the fruit cluster, the Indians wet and stretch 
it on the head to give it the desired fit (Appun 1871: 479; 
Wilbert 1963: 9). By pleating them and decorating them 
with bast ribbons the long-peaked caps sometimes acquire 
a bizarre appearance (PLates LXXV-LXXVII). They are 


[ 295 ] 


LXXIV 


PLATE 


[ 296 ] 


PuaTtE LXXV 


Hats made from spathe of Manicaria saccifera. 
Drawn by Hexica Ap1BI 


[ 297 ] 


PLATE LXXVI 


Warao Indian with hat made from spathe of Manicaria saccifera. 
(Courtesy P. T. Furst) 


[ 298 ] 


PLATE LXXVII 


[ 299 ] 


Warao Indian (on the left) with hat made from spathe of Manicaria saccifera. 


(Courtesy P. T. Furst) 


worn in many shapes and forms by natives, Creoles, and 
tourists in Mesoamerica (Standley and Steyermark 1958: 
273) and in Brazil (Wallace 1853: 70). I have not seen 
them used by the Warao as bags and wrapping cloth (ibid. ) 
or as loin cloths (Braun 1968: 111). 

The fruit. The fruits (yawihi aukwaha) serve the Warao 
as food and drink. One specimen of infructescence that I 
examined weighed 12 kg; the clear liquid inside the nuts 
accounted for approximately one-third, i.e., 3.75 liters (PLATE 
LXXVIII). The entire bunch was 87.5 cm. long and 25 cm. 
wide and was composed of 67 fruits: 22 one-seeded, 25 two- 
seeded, and 20 three-seeded ones.'° The single fruits mea- 
sured between 7 and 8 cm. in diameter, twins and triplets 
9.5 cm. All a Warao needs do to satisfy his thirst, whether 
in the jungle, along the coast, or in the field, is to cut just 
one bunch of temiche fruit and drink his fill. In times of 
non-potable water, the Indians take along a load of fruit 
clusters in their dugouts (Turrado Moreno 1945: 92). To 
get at the water, the Indian bites into the corky exocarp 
(nakoro ahoro) or cuts a hole in it and drinks the water 
from a calabash or from its natural cup. The fleshy homo- 
geneous endosperm of the as yet immature fruit (nohi) is 
greatly relished by the Indians; the fruit is opened with a 
blow of a heavy stick and the jelly-like substance scraped 
out with the thumbnail. Fully matured fruits, which re- 
semble miniature coconuts (ya umo) fall to the ground. 
They get buried in the detritus or are washed out to the 
rivers and the sea. In this state, they are hard and inedible, 
but find use as whorls in the manufacture of toy tops for 
boys (PLare LXXIX). 

The seedlings. Seedlings (emukohoko) are sought after 
by the Indians, young and old. If the germination results 
obtained by Braun (1968: 54) for cultivated Manicaria 
saccifera are any indication, the seeds of this palm germinate 
in four months, more or less. The Indians searah for seedlings 
that are developed enough to have grown their characteristic 
bifid eophylls up to 30 cm. above the ground.'' They pull 


[ 300 ] 


LXXVIII 


4 
vi 


PLATE 


LYaaTI AA SANNVHO[ hq ydvssojoyg 
me) 


OUIDS 


ayonijur pue oyyeds Suni 


cI 


Pa 


[ 301 ] 


PLATE LXXIX 


Top made of Manicaria seed. Drawn by Heica AprBI 


[ 302 ] 


them out and crack the hard shell of the seed with a heavy 
stick, exposing the white spongy haustorium. It has a mildly 
sweet taste, and as many as twenty at a time can be con- 
sumed without fear of digestive complications. Only children 
who eat primarily a diet of ocumo and fish are said to suffer 
occasionally from diarrhea. 

Ethnomedicine. Several parts are believed to have me- 
dicinal properties and are used as remedies against catarrh: 
that is, cough and fever; and against symptoms of other 
respiratory problems. An Indian woman prepared some 
medicines in my presence, so that I can verify the following 
recipes — if not attest to their effectiveness. 

To alleviate coughing and to repress fever, the water of 
Manicaria fruit is carefully strained through a cloth to 
eliminate impurities. The patient drinks a cup three times 
a day. 

Fever and cough are alleviated with a potion made of 
the green juices from the fresh eophylls of Mauritia mixed 
with Manicaria water. Added to the slimy chlorophyllous 
liquid is the urine of a child of opposite sex of the patient. 
The preparation is set aside for forty-eight hours and then 
applied to the patient’s body, especially his temples and 
forehead. The treatment is repeated three times a day. I 
found the remedy to have a definite cooling, hence soothing 
effect. Small children are washed in the liquid from head to 
foot to stop diarrhea accompanied by fever. Occasionally, 
they are also given a small quantity to imbibe. 

The anti-asthmatic and anti-catarrhal effect of the liquid 
endosperm of Manicaria has been mentioned by previous 
investigators. Since asthma does not seem to afflict the 
Warao, I could not verify this assertion, although all in- 
formants assured me that the fruit water of the palm 
facilitates breathing in congested patients. Most effective 
in this respect is a concoction made of grated Manicaria 
palmito mixed with fruit water. The remedy is imbibed. 

Indirect benefits. An indirect benefit accruing to the 
Indians from Manicaria relates to the fact that the ripe, 


[ 303 ] 


fallen fruit attract certain mammals like deer, peccary, paca, 
monkey, and agouti; turkey birds, like the pauji, are also 
fond of them. Traditionally, however, the Warao avoid the 
large species of mammal, like deer and peccary, and refrain 
from eating monkey altogether; but they like agouti, paca, 
and pauji. The agouti and the paca are ambushed at sunrise 
by the hunter ice hides in the vicinity of the palm. The 
birds are best shot in the early afternoon hours, when they 
give themselves away by the crashing noises that they pro- 
duce moving around among the leaves of the plant. 

I have mentioned the important protein supplement to 
the diet of these Indians provided by the rich supply of 
fatty larvae that are laid in Mauritia and in Manicaria by the 
palm borer 

amen of sago. In the summer of 1974, while collecting 
data on Warao ethnomedicine in the forests of the Orinoco 
Delta, one of my informants volunteered that, in addition to 
providing the Indians with a remedy against fever, catarrh, 
and diarrhea, Manicaria also produced a starch similar to 
that from the Mauritia. He added that the extraction of 
temiche sago had fallen out of practice, but that it had 
formerly hece common, Realizing that when a Warao speaks 
of palm starch he knows what he is talking about, I asked 
him to produce some temiche starch for me when I returned 
the following summer. What follows is a report on the 
process of sago recovery from Manicaria saccifera as wit- 
nessed by me in August 1975. PLatres LXXX-XCVIII 
illustrate the process. 

Throughout an area of dense pluvial forest scattered in- 
dividuals of Manicaria grew at distances from each other of 
4to6m. Their trunks (akabaho) were mostly erect and of 
varying heights, from 2 m. to 8 m. They were dark brown 
with very prominent circular leaf scars 2 to 3 cm. thick. 
Most of the palms seemed to be growing on small mounds 30 
to 50 cm. high, but their roots (ya ahokonamu) were not 
exposed, or only minimally so. 

After felling the palm, the Indian established how. far 


[ 304 | 


PLATE XX. 


Felling the Manicaria saccifera. Photograph by JOHANNES WILBERT 


[ 305 ] 


PLATE LXXXI 


[ 306 ] 


Removing the bark for sago extraction. 


Photograph by JoHANNES WILBERT 


[ 208 ] 


Making a trough from a section of Mauritia flexuosa. 


Photograph by JOHANNES WILBERT 


IIXXX'T ALVId 


down from the crown it contained sago. The testing was 
done by driving an axe into the trunk at various intervals. 
When the axe was withdrawn, with starch sticking to its 
blade, it signified contact with starchy pith. 

According to the Warao, Manicaria, unlike Mauritia, has 
no annual flowering and fruiting period and contains starch 
the year round. The specimen used for the experiment con- 
tained sago in the upper 3 m. of the 6 m. long and 30 cm. 
thick trunk. To obtain for me a unit measurement of the 
volume of sago in one palm, the Indian removed with his 
axe the bark of a 1 m. long section below the crown, exposing 
in the opening a beige to light brown fibrous interior, not 
pithy like Mauritia but somewhat ligneous. 

The Indian then stood on top of the trunk and shredded 
the pith with an adze or hoe (nahuru).’ The hoe is a 
composite tool which the Warao claim to have adopted, in 
remote times, from cannibalistic neighbors, called Siawani. 
It consists of three basic parts — blade, handle, and binding. 
The blade (nahuru ateho) is carved from the bark of a 
mature Mauritia and is 3 cm. thick. Its length varies between 
40 and 60 cm. according to the height of its user. The work- 
ing end (ahi) of the wooden blade is about 15 cm. wide and 
grooved to form a double cutting edge. Laterally, the blade 
is carved concavely and provided with notched shoulders 
(arokuaha) near the end opposite the cutting edge to facil- 
itate securing the blade to the cleft end of the handle. 

The handle (aka) is a round piece of wood 3 cm. thick. 
Any hard wood will do, and the length of the handle is 
roughly equal to the length of the blade. A cleft is made in 
one end, into which the non-cutting end of the blade is 
firmly wedged; the junction is lashed together with two-ply 
cordage made of Mauritia bast. A second string of this kind 
(ahutu) connects the blade with the handle like the crossbar 
of the letter A. To prevent this binding from slipping, two 
notches (iwiri) are made on the sides of the blade about 
20 cm. below the cutting edge (PLates LXXXIV-LXXXVI ). 

The Indian had made a new hoe the day before. As it 


[ 308 ] 


XXXII 


~ 
4 
wv) 


PLATI 


ro) 


INAGTI AA SANNVHO[ hq ydvss0j0Yg 


‘psonxayf DiyunDW 


fO YUNA oy} jo UOT{IIS wolf ogperul ysno.y pode 


ys 


A 


[ 309 ] 


PLATE LXXXIV 


Using a hoe to crush the starch. Photograph by JOHANNES WILBERT 


[ 310 ] 


PLATE LXXXV 


The opened Manicaria saccifera is lying on top of leaves to prevent 
the shredded pith from falling on the swampy soil. Close-up of the hoe 
used in the process of crushing the pith. 

Photograph by JOHANNES WILBERT 


[311 ] 


turned out, inexperience with extracting sago from Manicaria 
prompted him to make a cutting edge as wide as that used 
for extracting starch from Mauritia, which has a much larger 
trunk. The space between the hard bark on either side of 
the opening measured only 26 cm., much less than the trunk 
of a Mauritia customarily utilized for starch recovery. Thus, 
the edge proved to be too wide to be efficiently used. The 
Warao hoe bears close resemblance to sago hoes used in 
southeast New Guinea for the same purpose (Stohr, 1972, 
Fig. 35). 

After the pith was crushed, a woman washed it in a trough 
made from a piece of the trunk of a Mauritia 1.25 m. long. 
About 25 cm. at either end of the trough (canoa arua) was 
left untouched and the centre section excavated by means 
of an axe, so that a cross section was V-shaped (PLATE 
LXXXIII). 

After the men had placed the trough in a north/south 
direction (it must never be in the direction of the course of 
the sun), the woman drove four 1.20 m. long petiole sections 
of Mauritia (namoru) halfway into the water-logged ground 
next to the hollowed-out Manicaria (PLareE LXXXVIIT). 
Two uprights 75cm. apart stood on either side of the trough, 
and on top of these uprights the woman placed a dish-like 
strainer (bihi) made of strips of Ischnosiphon. Immediately 
below the strainer she positioned, at a steep angle, the fleshy 
end of a Mauritia leaf stem (wate buaka) for the purpose of 
collecting the washed pith below the strainer and channeling 
it smoothly into the trough without splashing. 

The woman collected the crushed pith from the trough, 
transferred it onto the strainer by means of a calabash 
(Cresentia Cujete), poured water over the pith, and began 
kneading it. From time to time, she scooped out some water 
from the trough and poured it over her hands and the pith. 
The water turned milky in the process, and the sago began 
settling on the bottom of the trough (PLAtes LXXXIX-— 
XCIII). 

The moment the woman started kneading the pith, she 


[312 | 


PrATE: LAXKV! 


Warao hoe used in palm starch extraction from both Mauritia 
flexuosa and Manicaria sacciferd. Drawn by HELGA ADIBI 


[ 313 ] 


[ FI8 | 


Trough with sieve resting on four uprights ready for the washing of Manicaria sac- 
cifera pith extracted from the palm in the foreground of the picture. Notice fleshy 
end of a Mauritia leaf stem below the strainer to channel the water into the trough 
without splashing. Photograph by JoHANNES WILBERT 


ALVId 


IAXXX'I 


[ cT¢ ] 


pl 


Trough filled with starch containing water after the woman finished washing the 
th. Photogrph by JOHANNES WILBERT 


ITAXXX'T S3LVTd 


[ 9T¢ ] 


Woman collecting shredded Manicaria saccifera pith into her calabash. 
Photograph by JOHANNES WILBERT 


LLVIg 


r. 
» 
e 


TXXX'T 


PLATE XC 


Calabash filled with shredded Manicaria saccifera pith resting on 
palm from which it has been extracted. 
Photograph by JOHANNES WILBERT 


[ 317 ] 


observed to me that it was rich in sago which, she said, one 
is able to determine by the viscid quality of the pith (PLATE 
XCIV). She observed, too, that it felt exactly like processing 
sago-rich Mauritia pith, except that she had to exert more 
pressure kneading the pith of Manicaria, because it was 
more ligneous than the other. Another difference between 
processing the two starches was that she had to wait longer 
for the Manicaria starch to settle at the bottom of the trough. 
She said that the resulting meal was of a lighter quality. 
After waiting for ten minutes in the shade of a windscreen 
(made of two temiche leaves stuck in the ground), the 
woman began to drain the water carefully by ladling it out 
of the trough with her calabash (PLate XCV). She then 
shaped the meal into a ball of light brown sago and, in doing 
so, proved conclusively that Manicaria saccifera must be 
counted among the sago-producing genera of the palms 
(PLATE XCVI). 

The total process of crushing, washing, and collecting the 
sago took approximately thirty working minutes. 

Upon completion of her work, the woman painstakingly 
washed her strainer to remove all particles from between 
the basketry strands (PLare XCVII). This act would pre- 
vent the strainer from rotting. Next, she pulled the four 
uprights out of the ground and tossed them to the side 
(PLate XCVIII). This would prevent malignant shamans 
from blowing on them, causing her arms to hurt. Finally, 
the man picked up the heavy trough and carried it a short 
distance away. This would make it more difficult for 
youngsters to poke inside with their machetes. People after 
mischief might do this to afflict the sago washer with ailing 
arms and shoulders. 

From the opening in the trunk, which was | m. long, the 
woman collected two calabashes packed to the brim with 
pith. The calabash used was 25 cm. long, 17.5 cm. wide, and 
12 cm. deep. From that much pith, the woman washed out 
750 gr. of sago. It had been ascertained that the trunk of the 
felled Manicaria contained starch within 3 m. of its upper 


[ 318 ] 


half, the total sago yield from that plant would amount to 
2.250 kg. I suggest adding at least another 750 gr. to account 
for the fact that the wide blade of the hoe prevented the 
Indian from extracting all of the available pith. A close 
approximation of the potential yield of the tested Manicaria 
is 3 kg. Even at that, Manicaria must be considered a low 
yield sago palm for the purpose to which the Warao put it: 
emergency food." 

In order to assess the nutritional value of Manicaria starch, 
I had the procured sample analyzed at laboratories in Cara- 
cas. The results of these tests are summarized in Table I. 


TABLE I 
Composition of Manicaria saccifera Starch 

pH (sol. 2% ) 5.75 
Humidity 63.51% 63.51% 
Fat * 0.55% 
Dextrose * 5.07% 
Protein °* 1.62% 
Starch * 4.57% 
Fiber ** 24.68% 

36.49% 36.49% 


100.00% 


* Calculation based on dry material 
°* By balance 


Ethnobotanical lore. The process of producing starch from 
Mauritia is a highly ritualized affair when done in prepara- 
tion for the annual harvest festival nahanamu or in connec- 
tion with any other propitiatory offering. The implements 
used in Manicaria starch extraction are also subject to certain 
taboos, but, for several reasons, Manicaria sago is considered 
less appropriate for these sacrificial purposes than the other. 

In the first place, the production yield of Mauritia is far 
greater (Heinen and Ruddle 1974). For instance, the amount 
of sago needed for a well prepared nahanamu festival varies 


[319 ] 


[ oz | 


Transporting a calabash full of shredded Maniacaria saccifera pith to 


the washing 
Photograph by JOHANNES WILBERT 


IOX @LV1g 


PLATE XCII 


[ 321 ] 


Woman pours the Manicaria saccifera pith onto the sieve. 


y JOHANNES WILBERT 


Photograph b 


from 500 to 1,500 kg., allowing from 2 kg. to 4 kg. per partici- 
pant. Families are engaged from six to ten weeks in its pro- 
duction. To produce this amount of sago from Manicaria 
would be almost impossible in terms of time and labor. The 
Warao insist, however, that Manicaria sago is acceptable to 
the Supernaturals and that it can be offered to them in pro- 
pitiation. 

A decisive factor in making palm starch acceptable to the 
gods is that, in its fresh state, it is practically odorless. Odors 
play an important role in communicating with the Super- 
naturals who accept as pleasing only the smell of tobacco 
smoke and that of carana (Protium heptaphyllum). Both 
types of sago available to the Warao fulfill this condition, but, 
besides relative abundance, Mauritia outdoes Manicaria 
on one other important score — it is colorless. When fresh, 
Mauritia sago is as white as plaster of Paris; whereas Mani- 
caria starch is beige to light brown. White is the color of 
predilection of the directional world gods who supposedly 
feel much less attracted by the color of Manicaria starch, 
which turns a dark brown several days after production. 

The color makes it much more appropriate for the Mani- 
caria to serve a secondary god, that of the dark Underworld. 
This spirit is known as Kanishabarao and is believed to dwell 
below the earth in company with his people, right next to 
the abode of Ya ahuba, the Temiche Master Snake. 

All palms, major trees, and most animals depend for their 
existence upon a master snake. The snake of Manicaria is a 
night spirit, appearing on earth only around midnight, when 
it comes to move softly the beautiful leaves of the palm. 
Thus, Manicaria pertains to the midnight sun as Mauritia 
belongs to the gods of the zenith and the world mountains 
at the cardinal and intercardinal points of the universe (Wil- 
bert 1973). 

This aspect recalls the day and/or night association of 
palms in other cultures. As Schultes (1974:7) pointed out, 
similar beliefs prevail among the Kuripako of Colombia in 
connection with Leopoldinia Piassaba. Fiber gatherers “are 
often bitten by poisonous snakes that infest the thick clumps 


[ 322 | 


of hanging fiber —a danger that probably underlies in part 
the natives’ belief that the evil spirit, the curupira, inhabits 
piassaba groves and wanders around at night.” The same 
curupira, we should add, is the central Master-of-Animals 
figure in South America who functions as the patron spirit 
of the trees and the forest (Zerries 1954: 18). Kanishabarao 
of the Warao is obviously a cousin of the Amazonian curupira. 
Furthermore, there exists also a sun-palm relationship among 
the Yukuna Indians of Colombia, whose kai-ya-tee festival 
resembles the annual sago festival, nahanamu, of the Warao, 
and “basically celebrates the harvest of the pupunha palm 
which was given to the Yukuna people by the ‘Sister of the 
Sun’ as one of their major cultivated foods” (Schultes 1974: 
16). 

That the Manicaria belongs to the midnight sun rather than 
to the day sun becomes apparent in the belief that, while its 
sago may be unsuitable for a nahanmu festival in honor of 
the cardinal gods, it is, nevertheless, used for this purpose by 
Kanishabarao and his people of the underworld. Annually 
they prepare a feast of temiche sago, and that is why the 
Indians may come across hollowed-out trunks of the palm in 
the forest. Kanishabarao and his people also eat the nuts and 
all the parts of the palm that humans enjoy. 

Finally, Manicaria sago is believed to be extracted by the 
monkey (naku) for purposes of celebrating an animal na- 
hanamu palm festival. Monkeys are supposedly very fond of 
the starch and are believed to be yet other ‘ ‘people” that 
hollow out the palms one happens across in the forest. 
Instead of fish or crab, monkeys are said to prefer large 
spiders (abunamoko) with their starch. 

The custom of recovering palm starch from Manicaria 
is essentially a feature of the past with the Warao. It came 
to my attention when an elderly informant remembered hear- 
ing his uncle and other elders send workers out to prepare 
Manicaria sago. It was done occasionally, he said, when the 
people were away from the Mauritia groves fishing on a ma- 
jor river. To supplement their diet, they relied on Manicaria 
sago which was always close at hand in the Intermediate 


[ 323 ] 


PLATE XCIII 


Beginning the process of washing starch out of the shredded pith 
of the Manicaria saccifera. Highly contaminated water is taken di- 


rectly from the swamp. 
Photograph by JOHANNES WILBERT 


PLATE ACV 


ams 


Woman kneading the pith with both hands, 


Photograph by JoHANNES WILBERT 


[ 9z¢ | 


After the starch has settled down at the bottom of the V-shaped trough the woman 


starts ladling out the water. Photograph by JOHANNES WILBERT 


AQX SLVIg 


PLATE XCVI 


Starch is scooped with hands from the bottom of the trough into a 
calabash. Photograph by JOHANNES WILBERT 


[ 327 ] 


LHAGTI AA SANNVHO[ fig ydvisojoyg ‘TeUa}eul Aayseq ay} Jo 
BuQ}OL ploae 0} Ay[Nyorvo AIOA JOUTeIYS OY} seyseM ULWIOM Yd ayy SuIssad0id J9z;y 


ee 


[ 328 | 


PLATE XCVIII 


Woman dismantles the washing stand after extracting enough starch. 
Photograph by JOHANNES WILBERT 


[ 329 ] 


zone of the Delta. The wife of my informant, who comes 
from a different region than her husband, said she also dis- 
tinctly remembered having seen people collect and eat 
temiche starch. Those who still know about it are in their 
fifties and older. The younger people in the village had never 
seen or even heard about it. 

One need not invoke prophetic insight to predict the dis- 
appearance, in the near future, of palm-starch production 
from all of South America. The majority of southern tribes 
referred to in the introduction have already either given it 
up or are so reduced in number that the practice of starch 
recovery will soon come to an end, Among the Warao, even 
Mauritia extraction is declining rapidly and will soon be- 
come a rarity. Yet, in South America, as elsewhere, recovery 
of starch from different genera of palms represents the sur- 
vival of a once more common tradition that reaches back 
into the remote, prehistoric past. It survived the Neo-Indian 
revolution of subsistence agriculture only among a small 
number of marginal tribes, but it is giving way increasingly to 
agricultural staples even among them. In any event, for the 
Warao this form of traditional aboriculture provided a mea- 
sure of economic stability not unlike that achieved by other 
indigenous societies through agriculture. The entire man- 
palm relationship as it developed among these people over 
many centuries can be fully understood only in the context of 
Manicaria, Mauritia, and Euterpe ethnobotany. Realizing 
this and going a little beyond the scope and evidence of the 
present paper, may I suggest in concluding that economic 
stability is not the only benefit that the Indians have derived 
from gathering around these regal plants. As their ethnobo- 
tanical lore reveals, the palms have nurtured among the 
Warao an exquisite partnership between man and nature — 
a symbiosis that, in addition to a viable socio-economic blue- 
print, generated an ideological matrix that gave meaning to 
the world and purpose to life. Surely, achievements of this 
sort must rank among the finest that mankind has made 
anywhere. 


[ 330 ] 


pea 


10. 


11. 


12. 


ENDNOTES 


. Cf. Lévi-Strauss 1950: 469-472; Fuerst 1970: 114-122. 
. The only previous mention that I could find was Barral (1949: 


150), who said: “Pero el “aru” propiamente tal, el “arw” guaraino 
[Warao], es la torta hecha con la fécula extraida de la “Mauritia 
flexuosa”, 0 también del temiche (Phitalephas temiche), no tan 
ponderada como la de moriche, pero tan real.” 

Among the secondary palms, the authors (ibid., 116) list: Acro- 
comia, Caryota, Coelococcus, Corphyra, Eugeissona, Phoenix, and 
Pholidocarpus. 


. Iam using the so called ethnographic present, although the prac- 


tice of palm-starch production may very well have been discon- 
tinued now among some of the tribes of southeastern South Amer- 
ica. 

Gumilla 1791, 1: 145; Schomburgk 1848: 49; Turrado Moreno 
1945: 73-83; Suarez 1966; Wilbert 1972: 81-82: Heinen and 
Ruddle 1974. 


. Cf. Dahlgren 1936: 202 and references since then (1936): Bailey 


1943: 392-393 Dugand 1940: 43; McCurrach 1960: 129-131; 
Wessels Boer 1965a, b; Braun 1968: 111. 


. If it is any consolation to botanists, Manicaria is ethnobotanically 


all but unknown; the snatches of information, frequently repeated, 
are restricted largely to the usefulness of the leaves for thatch 
and of the spathe for “monkey-caps.” In estimating migration and 
settlement patterns, for instance, archaeologists and ethnologists 
may have underestimated the effect of palm exploitation on the 
relative stability of autochthonous peoples in South America and 
elsewhere. I would like to thank Mr. August Braun and Drs. 
Harold E. Moore, Jr., Richard Evans Schultes, and Julian A. 
Steyermark for advice and assistance with the botanical aspects 
of this paper. 

Von Spix and von Martius (1823-1831; 3, 989) observed the same 
custom practiced on Marajo Island in the mouth of the Amazon. 
Lat. manica = sleeve; saccifera = sack-bearing. 

The ratio of single to multilobed fruits seems to vary markedly. 
In a second sample examined, I counted a total of 72 fruits with 
42 one-seeded, 27 two-seeded, and three 3-seeded ones. 

The apical incision in Manicaria eophylls results in two opposite 
terminal leaflets, each segment with an acute apex and smooth 
margins. 

The hoe and all the other tools and techniques employed in the 
recovery of Manicaria starch are the same ones that the Warao 
use for the extraction of sago from Mauritia. 


[ 331 ] 


13. I am taking the Indians’ word for the fact that Manicaria has no 
annual flowering period and that the palm carries starch the year 
round. If incorrect, the result of a single test ought not to be 
generalized, since it is known that the starch content in Mauritia 
of the Orinoco Delta, for instance, may vary seasonally from 2 kg. 
to 60 kg. per palm (Heinen and Ruddle 1974: Peae 


[ 332 | 


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