BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY
VOLUME XIV
OAKES AMES MEMORIAL VOLUME
BOTANICAL MUSEUM
CAMBRIDGE, MASSACHUSETTS
1949-1951
PRINTED AND PUBLISHED AT THE
BOTANICAL MUSEUM
CAMBRIDGE, MASSACHUSETTS
From a painting by Blanche Ames, 1946
OAKES AMES
1874-1950
YEAR has now elapsed since ProfessorOakes Ames,
who served the Botanical Museum as its principal
administrative officer from 1928 to 1945, first as Curator,
later as Supervisor, and finally as Director, passed away
on April 28, 1950. In the intervening months the mem-
bers of the staff of the Botanical Museum have been
engaged in completing this volume of the Botanical
Museum Leaflets which, soon after his death, it was de-
cided to dedicate to his memory.
The choice of the Leaflets as the medium for a mem-
orial volume was a quite natural one since Professor
Ames himself founded the Leaflets and since they rep-
resent not only his diverse botanical interests, but also
his insistence upon high quality in paper, printing, com-
position and illustration. Professor Ames deplored the
trend toward cheaper publications. ‘‘A scholarly scientific
paper,” he said, ‘‘is a jewel worthy of a proper setting. ”’
The entire staff of the Botanical Museum has shared
in the production of this volume, and the papers which
it contains represent quite accurately the extraordinary
diversity of Professor Ames’ botanical interests. Of the
fourteen papers, ten are concerned with taxonomy and
three of these with the taxonomy of the orchids, Professor
Ames’ lifelong special interest. His second professional
love, economic botany, is represented by eight papers
of which five are also taxonomic and three are in ethno-
botany, a field which the Botanical Museum, at Professor
Ames’ instigation, has fostered in recent years. Paleo-
[ vii ]
botany, a traditional interest of the Museum and one
which Professor Ames was instrumental in reviving, is
represented by one significant paper. Professor Ames’
sustained interest in the course in Economic Botany,
which he taught for many years, finds representation in
“The Story of Pan Chewing in India’ which was sub-
mitted as a term paper by a student enrolled in the
course.
Only one facet of Professor Ames’ botanical career
fails to find expression in this volume; his broad philo-
sophical outlook upon the Orchidaceae, a plant family
to whose study the major part of his life was devoted.
Happily this interest had previously been treated in
‘*Orchids in Retrospect,’” a collection of his essays on
the Orchidaceae which was published in 1948 by his
colleagues to commemorate the fiftieth anniversary of
his service to Harvard University. This book also con-
tains a biographical sketch.
With deep respect for the memory of Oakes Ames,
with constant admiration for the standards of excellence
for which he stood, and with a firm purpose of perpet-
uating the sound traditions which he established, the
members of the staff of the Botanical Museum dedicate
this memorial volume.
Cambridge, Massachusetts
April 28, 1951
[ viii ]
TABLE OF CONTENTS
NuMBER 1 (June 27, 1949)
Degradation of Plant Remains in Organic Sedi-
ments
By Etso S. BarRGHOORN .
NuMBER 2 (October 26, 1949)
Plantae Colombianae XII
By Ricuarp Evans SCHULTES
NuMBER 8 (December 7, 1949)
Nomenclatural Notes and New Concepts of Tropi-
cal American Orchids
By CHARLES SCHWEINFURTH .
Noumse_er 4 (February 6, 1950)
Studies in the Genus Hevea III
By Ricuarp Evans ScHULTES
The Identity of Ucuqui
By JoAo Murca PirEs aND RicHarp Evans
SCHULTES
The Correct Name of the Yaupon
By Ricuarp Evans SCHULTES
A New Variety of [lex vomitoria from Southern
Mexico
By A. J. SHarp
[ ix ]
21
49
87
97
107
Number 5 (March 20, 1950)
Plantae Austro-Americanae VI
By Ricuarp Evans SCHULTES
Numser 6 (November 6, 1950)
Notes on Peruvian Cinchonas I
By W. H. Hopce
Number 7 (December 11, 1950)
Prehistoric Plant Remains from Bat Cave
By C. E. Smita, Jr.
NuMBER 8 (January 15, 1951)
The Story of Pan Chewing in India
By M. Gowpa
NuMBER 9 (March 138, 1951)
New Orchids from Africa
By V.S. SUuMMERHAYES .
A Revision of the Genus Angraecopsis
By V. S. SUMMERHAYES
NuMBER 10 (April 18, 1951)
Whence Came Maize to Asia?
By Paut C. MANGELSDORF AND Doucutas L.
OLIVER
[x ]
109
137
157
181
215
240
263
INDEX OF ILLUSTRATIONS
PLATE
pamece Cacecnt.c 2 « fa « 5% > OSG MEV
Ambelania zschokkeiformis Marcg. . . . . . XXX
Brassavola ovaliformis C. Schweinf. . . . . . XVII
Cecropia porvenirensis Cuatr. ........ VII
Cerro del Castillo, Rio Apaporis, Colombia. . . . X
Cinchona glandulifera R. & P. . . . . . XX XVIII
Cinchona Pahudiana Howard. . . . . . XXXIX
Cinchona parabolica Pav. . ......... XL
Combretum karijonorum FP. EH. Schultes . . XXXVI
Cranichis ciliilabia C. Schweinf. . . . . . . . . XII
Degraded wood ......... LILIUO,IV
Discussion of the black-drink, in Catesby’s
**Natural History of Carolina, Florida and
the Bahama Islands” ......... XXXVI
Distribution of Areca Catechu Z. in eahivaiion
GD) cs aces sc SS ee A ew De EVE
Distribution of the betel-chewing habit (Map) XLVI
Earliest publication of the name Stphonia
brasiliensis, in Adr. de Jussieu’s ‘‘De
Euphorbiacearum generibus” . . . .. . XVIII
[ xi ]
Epidendrum flexuosissimum C. Schweinf. . . XIV
Epidendrum pajitense C. Schweinf. . . . . . . XV
Epidendrum strictiforme C. Schweinf. . . . . XVI
First Kuropean account of the black-drink,
in de Vaca’s ‘‘Relacién y commentarios’””» . XXIV
POSSI WOOK, 4.6% oe be ee eR Oe
Herrania Camargoana R. FE. Schultes X XIX, XXXII
Herrania kanukuensis R. H. Schultes . . . XNXXIII
Herrania kofanorum R. E. Schultes XX VIII,X XXIV
Herrania Mariae (Mart.) Decaisne ex Goudot
var. putumayonis R. Hf. Schultes . . . . . XXX
Herrania nycterodendron R. EH. Schultes . . XXXV
Herrania pulcherrima Goudot var. pacifica
R. E. Schultes ........... . XXVIII
Hevea brasiliensis (Willd. ex Adr. de Juss.)
DTU MRATER 6c i i we we ea we Ee Ke
Ilex vomitoria (Solander in) Mit. . 2. . 2. XXV
Lindakeria nitida Aullip & Schultes. . . 2... . IX
hbue glomerata Killip & Schultes. . 2... . XI
Normal wood. .......... 2.2... 1V, V
Octomeria pygmaea C. Schweinf. . . . . . . XIII
Parascheelia anchistropetala Dugand. . . . XXVII
{ xii ]
Piper Betle LL. ........2.4..+.,. MLVII
Plains of San Augustin, New Mexico, near
Pee | CS a ke ee eg eee
Plant remains from Bat Cave. . . . XLII-XLIV
Pouteria Ucuqui Pires & Schultes . . . XX-XXIII
Rapata modesta Maguire ........ =XXXI
Rhytidanthera mellifera R. H. Schultes. . . VIII, X
Roots isolated from post-glacial peat. . . . .. VI
Sandia Valley and Ramospata Ridge (Dept.
of Puno) Peru ........... ~XXXVII
Solanum quitoense Lam. ...........3X
Types of beeda used in pan-chewing . . . . . XLVI
Vochysia lomatophylla Standl. . ...... =. VII
ZeaMaysh. ............ .XLVIII
[ xii |
INDEX
TO GENERA AND SPECIES
ACACIA
Catechu Willd., 195,209
ACIOTIS
aequatorialis Cogn., 40
polystachys (Bonpl.) Tr., 40
purpurascens (Aubl.) Tr., 40
adike, 185,206
AERANGIS, 240
AERANTHUS
trifurcus Reichb.f., 252,253
AEROBION
parviflorum Spreng., 255
agase, 210
AGERIA
Cassena (Michx.) Raf., 98
atoho-ayo-o, 40
ajuo-wo, 240
allu, 210
AMARANTHACEAR, 172
AMARANTHUS
blitoides S. Wats., 172,179
hybridus L., 172,179
Powellii, S. Wats., 172,175,
179
sp:,-172,179
amarilla, 154
AMBELANIA
cuneata Muell.-Arg., 135
zschokkeiformis Markg., 135
amber, 197
amli, 209
AMOMUM
subulatum Roxb., 206
ANACARDIUM
occidentale L., 212
ANETHUM
graveolens L., 208
ANGRAECOPSIS, 240,241,
242
amaniensis Summerh., 246,
260,261
Boutoni (Reichb.f.) Perr. de la
Bathie, 261
breviloba Summerh., 244,246,
261
sect. Cardiochilus, 243,258
sect. Coenadenium, 244,259
comorensis Summerh., 252
dolabriformis (Rolfe) Schltr. ,
245,252,253,255
elliptica Summerh., 245,250
sect. Eu-angraecopsis, 243,
246
Jalcata (Thunb. ) Schltr., 240,
241
gracillima (Rolfe) Summerh.,
244,248, 249,250
holochila Summerh., 246,260
ischnopus (Schltr. ) Schltr.,
24.0, 245,247,252
macrophylla Summerh., 245,
247,249
[ xv ]
occidentalis Schltr., 254
parviflora (Thou.) Schltr.,
240,246,255, 256
Pobeguinii (Finet) Perr. de la
Bathie, 246,257
pusilla Summerh., 244,246,
258
tenerrima Krénzl., 240,248,
244,247,248
tenuicalear Summerh., 260
Thouarsii Perr. de la Bathie,
252,253
tridens (Lindl. ) Schltr., 245,
252,254
trifurea (Reichb f.) Schltr.,
245,250,252,257
ANGRAECUM
amaniense Krinzl., 247
Boutoni Reichb.f., 261
dolabriforme Engl., 254
ischnopus Schltr., 247
occidentale Rolfe, 254
parviflorum 'Thouars., 255
tenerrimum Schltr., 247
tridens Lindl., 254
APLOPAPPUS
gracilis (Nutt.) 4.Gray, 174,
179
APOCYNACEAE, 43,135
AQUILARIA
Agallocha Roxb., 207
ARACEAE, 22
arali, 210
ARECA
Catechu L., 185,186,196,
206
var. silvatica Becc., 187
arrow-poisons, 30,41,42,43,120
ARTEMISIA
spp., 174,179
ARTOCARPUS
altilis (Park.) Fosb., 212
heterophyllus Lam., 212
ASCLEPIADACEAE, 173
ASCLEPIAS
sp., 173,179
Assamese maize, 265
asvattha, 210
aswat, 210
ATRIPLEX
canescens (Pursh) Nutt., 158,
171,179
BACTRIS
balanophora Spruce, 110
bale, 211
battalu adike, 193
beeda, 190,200,201,202
be-ee-cha, 111
BETA
vulgaris L., 209
Betel chewing, 181,190
Betel leaf, 181,182,191,195,
204,206
Betel nuts, 181,182,184,185,
191, 192,208,206
Betel palm, 185,186,187,190,
205
Betel pepper, 188,206
Betel vine, 188,189
betre, 188
boia, 136
BomBacackak, 30
{ xvi |
BoraGinaceak, 158
BOSWELLIA
serrata Rovb., 211
BOUTELOUA
curtipendula (Michr.) Torr.,
159
gracilis (HBK.) Lag. ex
Steud., 168,178
hirsuta Lag., 158,159,168,
175,179
BRASSAVOLA
nodosa (L.) Lindl., 61
ovaliformis C.Schweinf., 60
Perrinii Lindl., 61
BROTOBROMA
aspera Karst. & Triana., 130
BULBOPHYLLUM
encephalodes Summerh., 228
lupulinum Lind/,, 230,232
oreonastes Feichb f., 228
porphyrostachys Summerh.,
230
unifoliatum DeWildem., 228
buntata, 285
cacao de jacaré, 124
cacao de monte, 129,132
cacao de macaco, 124
cacao gacaré, 123
cacao silvestre, 128,129,130
CacTAcEAk, 173
CALAMAGROSTIS
inexpansa 4.Gray, 167,178
capavira, 116
CARUM
Bulbocastanum Koch, 208
Carvi L., 208
CASINE
yapon Bartr., 98
CASSINE
? amulosa Raf., 98
caroliniana Lam, 98
corymbosa Mill., 105
Paragua Mill, 98,100,101,
103
Peragua L. (1762), 98
Peragua L. (1771), 101
? ramulosa Raf., 98
vera Floridanorum.... Pluk.,
99,100
vomitoria Swanton, 99
yaupon Gatschet, 99
cascarilla crespilla, 153,155
cascarilla crespilla chica, 150
cascarilla crespilla con hajas
rugosas, 155
cascarilla del pajonal, 147
cascarilla delgada, 147
cascarilla negra, 147
cascarilla negrilla, 147
CASTILLEJA
minor 4.Gray, 158
catechu, 185,195,209
CECROPIA
discolor Cuatr., 25
latiloba Miqg., 118
marginalis Cuatr., 25,26
mocoana Cuair., 24
porvenirensis Cuatr., 25
telealbida Cuatr., 118
telenivea Cuatr., 118
tolimensis Cuatr., 118
chakkota, 212
[ xvii ]
CHAMAEANGIS, 247
Hariotiana (Kriinzl.) Schltr.,
261
Pobeguinit Schltr., 257
CHAMAEDOREA
integrifolia (T'rail) Dammer,
110
lanceolata (Ruiz & Pav.)
Kunth, 110
champaca, 212
chandan, 210
cha-te-ré, 131
cheape, 212
Cuenopopiaceak, 160,171,179
CHENOPODIUM
sp., 171,179
chimbillo, 27
CHLORANTHACEAE, 117
chongaru, 193,196
chua, 197
churu-adike, 194
CINCHONA
Calisaya Wedd., 139,141
carabayensis Wedd., 139,140,
141,142,147, 149
Delessertiana Standl., 150,151
glandulifera Ruiz & Pav.,
143, 144,145,150
Josephiana Wedd., 141,145
micrantha Ruiz & Pav., 143
Mutisii Lamb.
var. B, 151
var. crispa Wedd., 151
officinalis, 147,154
Pahudiana Howard, 141,142,
147,148,149
parabolica Pav. ex Howard,
150,151,152,153,154
pubescens sensu lat., 188,145,
149,154
rugosa Pay. ex Howard, 151,
153
var. crispa Wedd., 151
succirubra Pav. ex Klotzsch,
138
CINNAMOMUM
Camphora (L.) Nees &
Eberm., 207
Cassia (Nees) Nees ex Bl., 207
zeylanicum Breyn, 207
CITRUS
aurantifolia (Chr.) Swgl., 211
grandis (L.) Osbeck., 212
Limon (L.) Burm f., 212
medica L., 212
sinensis (L.) Osbeck, 212
CLIBADIUM
asperum (Aubl.) DC., 46
spp., 47
COCOS
nucifera L., 207,211
colorada, 154
ComMBRETACEAE, 39,134
COMBRETUM
karijonorum R. E.Schultes,
134
laxum Jacq., 39,40
rotundifolium L. C. Rich., 135
ComposITak, 46,158,160,161,
174,179
coqui, 90
CORIANDRUM
sativum L., 208
[ xvii |
COUSSAPOA
intermedia Mart. ex Miq., 119
magnifolia Trécul, 119
CRANICHIS
ciliilabia C.Schweinf., 49
crespilla, 153
CROCUS
sativus L., 207
croker, 285
CUCUMIS
Melo L., 209
CUCURBITA
foetidissima HBK., 174,179
Pepo L., 161,162,164,177,
178
CucurBITACEAR, 174
CUMINUM
Cyminum L., 208
CUNURIA
crassipes Muell.-Arg., 54
curare, 41,42,43,120
CURCUMA
longa L., 212
cutch, 198,195
CYPERACEAR, 22,169
CYRTOCHILUM
aemulum Kranzl., 73
falcipetalum Krinzl., 68
hastiferum Krianzl., 70
Incarum Krinzl., 70
monachicum, 69
mystacinum Lindl., 66
Pavonii Krinzl., 68
superbiens Krinzl., 73
undulatum HBK., 73
ventilabrum Kranzl., 73
Weberbauerianum Krinzl., 64
CYTOPHAGA, 12
dalchini, 207
dalimbre, 212
darchini, 207
dhanya, 208
DIAPHANANTHE, 247
dodda-jirige, 208
DRYOBALANOPS
aromatica Gaertn f., 207
ECCLINUSA
Balata Ducke, 91
sanguinolenta Pierre, 91
echenique, 141
ee-ru-che, 42,43,44
ee-sb-pe-ke, 123
EGGELINGIA, 235
clavata Summerh,, 238
ligulifolia Summerh., 236,238,
239
elakki, 206
ELETTARIA
Cardamomum (L.) Maton.,
206
ELYMUS
salina M. EF. Jones, 167,178
? EMETILA
ramulosa Raf., 99
EPIDENDRUM
exasperatum Feichb,f., 58
flexuosissimum C.Schweinf.,
ao
pajitense C.Schweinf., 56
sarcodes Lindl., 60
Schumannianum Schltr., 58
[ xix ]
strictiforme C.Schweinf., 58
verrucosum Sw,
var. myrianthum
(Lindl.) Ames & Correll,
58
EPIDORCHIS
parviflora O. Ktze, 255
tridens O.Ktze., 254
ERYTHRINA
indica Lam., 188,211
spp., 189
EUPATORIUM
sp., 174,179
EUPHORBIA
penicillata (Millsp.) R.E.
Schultes, 29
Tirucalli L., 211
EupuHorBIAcEAk, 29
Facaceak, 170
FALLUGIA
paradoxa (D. Don) Endl., 158
See-see-pa-chu, 43
FESTUCA
Kingii (8S. Wats.) Cassidy,
167,178
FICUS
religiosa L., 193,210
FIMBRISTYLIS
miliacea (L.) Vahl, 22
Jish-poison, 47
FLACOURTIACEAE, 36
FOENICULUM
vulgare Mill,, 208
FORSTIERA
sp., 158
gandham, 210
[ xx ]
ganna, 209
GEONOMA
interrupta (Ruiz & Pav.)
Mart., 111
laxiflora Mart., 111
pycnostachys Mart., 110
geru, 212
GODOYA, 34
godumbi, 212
GONGYLOLEPIS
maroana Badillo, 47
GOSSYPIUM
arboreum L,, 286
barbadense L., 287
Darwinii Watt, 287
hirsutum L,, 287
Raimondii Ulbr., 286
tomentosum Nutt., 287
GRAMINEAE, 167
granisillo, 117
granisillo del grande, 118
granisillo pequeno, 117
guaco, 47
guamo machete, 27
guava machete, 27
gundi, 197,203
gurvaca, 206
HABENARIA
§ Diphyllae, 217
egregia Summerh., 217
falciloba Summerh., 215
Keayi Summerh., 217
Lecardii Krénzl., 219
§ Multipartitae, 215
pilosa Schltr., 217,219
halasu, 212
harisina, 212
he-de-kd-pe, 120
HEDEOMA
Drummondii Benth., 158
HEDYOSMUM
toxicum Cuatr., 117
translucidum Cuatr., 117
HEISTERIA
cyanocarpa Poepp. & Endl.,
119
eurycarpa Stand/l., 120
HELIANTHUS
annuus L., 164,175,178
HELICONIA
lingulata Ruiz & Pav., 116
psittacorum L,f., 116
Schumanniana Loesn., 116
stricta Huber, 117
HELIOTROPIUM
curassavicum L,, 158
HERRANIA
albiflora Goud., 131
aspera (Karst. & Triana)
Karst., 130
balaénsis Preuss, 128
Camargoana R. EF. Schultes, 120
Dugandii, 128
kanukuensis R. FE. Schultes, 126
kofanorum R.E.Schultes, 126
128
lemniscata (Schomb.) R.E.
Schultes, 125,126
laciniifolia Goud. ex Triana &
Planch. apud Garcia- Barriga
125
Mariae (Mart.) Deen. ex
Goud., 125,129
var, putumayonis R, F.
Schultes, 129
nitida (Poepp.) R.E.Schultes,
130
var. aspera (Karst. &
Triana) R. E.Schultes,130
forma sphenophylla R. E.
Schultes, 131
var. sphenophylla R.E.
Schultes, 131
nycterodendron, 129
pulcherrima Goud., 131,132
var. pacifica R. EF. Schultes,
131
purpurea ( Pitt.) R.E.
Schultes, 132,133
hess-pda-chu, 41
HEVEA
brasiliensis (HBK.) Muell.-
Arg., 85
brasiliensis (Willd. ex A. Juss.)
Muell.-Arg., 79,80,85
var. acreana Ule, 86
var. angustifolia Ule, 86
mut. Granthami Bartlett,86
var. janeirensis (Muell.-
Arg.) Pax, 86
var. latifolia Ule, 86
forma Randiana (Huber)
Ducke, 86
var. Randiana (Huber. )
Pax, 86
forma subconcolor Ducke, 86
var. subconcolor Ducke, 86
var. stylosa Huber, 86
forma typica Ducke, 86
Granthami Bartlett, 86
Janeirensis Muell.-Arg., 85
Kunthiana Ule, 80
Randiana Huber, 86
Siebert Warb., 80,81,85
[ xxl ]
HIEROPHYLLUS
Cassine (Walt.) Raf., 98
HIMATANTHUS
articulata (Vahl) Woodson, 43
bracteata (4.DC.) Woodson,
44
hogesoppu, 209
hunise, 209
ILEX
caroliniana (Lam.) Loes.,
97,99,105
Cassena Michx., 98
Cassine L., 103
Cassine Walt., 98,103
Cassine B L., 98,100
floridana Lam., 98
ligustrina Jacq., 98
Peragua (L.) Trel., 99
religiosa Bartr., 98
vomitoria [Soland. in] Ait.,
97,98, 100, 104,105,107
var. chiapensis A. J.Sharp,
107
var. Yawkeyit Tarbox, 99
INGA
macrophylla H. & B. ex
Willd., 27
setifera DC., 27
jaiphal, 207
jambul, 209
jatri, 207
jayanti, 210
Jharada, 196,203
Jjiluga, 210
Jira, 208
Jirige, 208
JUGLANDACEAE, 170
JUGLANS
major (Torr.) Heller, 170,
175,178,179
JUNIPERUS
pachyphloea Torr., 165,175,
178
spp., 159,166,178
kabbu, 209
kachu, 185,209
kadapan, 197
kaddipudi, 197
ké-he-pa, 89,90,91
kala zerah, 208
karyirige, 208
karpura, 207
kath, 193,195,196,209
kau-ré, 27
kela, 211
kempugandha chekke, 210
kesar, 207
kesari, 207
khaina, 196
ko-kee-ot-chu, 128,129
kolu kalli, 211
konda mavu, 211
kothamburi, 208
kumkumkesari, 207
ku-na-pe, 47
kun-kun-ni-ai, 30
ku-see-ye-hé-pa, 42
LaBiATAE, 158,173
LAPPULA
[ xxii |
Redowskii (Hornem.) Greene,
158
lavang, 206
lavanga-churu, 194
LEANDRA
divaricata (Naud.) Cogn., 40
le-che-mah, 135
Leeuminosak, 27,158,173
LEMNA
valdiviana Philippi, 23
LeMNACEAR, 23
LEPIDOCARYUM
tenue Mart., 111
Litiaceak, 169
lime, 197
LINDAKERIA
maynensis, 37
nitida Killip & R.E.Schultes,
36
LIPARIS
Deistelii Summerh., 226,228
odontochilos Summerh., 226
LISTROSTACHYS
parviflora S.Moore, 255
tridens Rehb.f., 254
trifurca Finet, 252
LogGanraceak, 41
laja, 154
lulo, 44,45
LYCIUM
sp., 158,173,179
MAHUREA
tomentosa Ducke, 133
maiz indio, 273
MALAXIS
Chevalieri Summerh., 226
Hirschbergii Summerh., 225
katangensis Summerh., 221
S Katochilos, 221,223,224
Maclaudii (Finet) Summerh.,
225,226
pygmaea Summerh., 221,223
MAMMILLARIA
sp., 159
MANDEVILLA
nerioides Woodson, 44
MANGIFERA
indica L., 211
MANIHOT
esculenta Crantz, 213
MANILKARA
spp., 92
maraganesu, 213
ma-sha-kve, 45
masala, 198,202
mavu, 211
MAYNA
glomerata Killip & R.E.
Schultes, 38
longifolia Poepp. & Endl., 39
MELASTOMACEAE, 40
MENISPERMACEAR, 120
MENTZELIA
pumila (Nutt.) T. & G., 158
MESOSPINIDIUM
Wallisii Reichb.f., 67
MICHELIA
Champaca L., 212
misreys, 208
mita zerah, 208
[ xxiii |
MONARDA
sp., 178,179
Moracekagk, 24,118
MORINGA
oleifera Lam., 210
MUHLENBERGIA
rigens (Benth.) Hitche., 168,
178
muhuri, 208
mun-tai-ru-chee, 42
murphy, 285
MUSA
paradisiaca L., 211
subsp. sapientum (L.)
O. Ktze., 211
Musaceak, 116
mu-se-nda, 130
MYRISTICA
fragrans Houtt., 206
MYSTACIDIUM, 242
dolabriforme Rolfe, 253
gracillimum Rolfe, 248
pedunculatum Rolfe, 255,256
Thouarsii Finet, 252
tridens Rolfe, 254
trifurcum Durand & Schinz,
252
naranjillo, 45
nariyal, 207
NEOFINETIA, 241
nerale, 209
NICOTIANA
rustica L., 209
Tabacum, L., 209
NIGELLA
sativa L., 208
nimbu, 211
nugge, 210
Ocunackak, 82
OCTOMERIA
minuta Cogn., 54
pygmaea C. Schweinf, 53
ODONTOGLOSSUM
angustatum Lindl. 1837, 62,
638
angustatum Lindl. 1846, 62
aureo-purpureum Reichbf.,
63,64
bellum Schltr., 62
brevifolium Lindl., 64,65
var. Weberbauerianum
(Kransl.) C. Schweinf. , 64
compactum Reichb.f., 63
flavescens Rolfe, 65
Koehleri Schltr., 63,64
Loesenerianum Schltr., 62,63
longifolium Lindl., 67
mystacinum ( Lindl.) Lindl.,
66
obscurum C.Schweinf., 67
retusum Lindl,, 65
rigidum Lindl., 66
tetraplasium Reichb.f., 62
Wallisii Lind. & Reichb.f., 67
Weberbauerianum Schltr., 64
Wyattianum G. Wils., 67
OECEOCLADES
parviflora Lindl., 255
OLacackAk, 119
ONCIDIUM
aemulum Reichb.f. & Warsc.,
1%,19
bryolophotum Reichb.f., 69,70
chrysopyramis Reichb.f. &
Warsc., 72
[ xxiv ]
Davisii Reichb.f., 68,69
falcipetalum Lindl., 67,68
hastiferum Reichb.f. & Warsce.
70
heteranthum Poepp. & Endl.,
69,70
incarum (Krdnzl.) C. Schweinf.
70
inferlobum Hort., 72
inops Cogn. & Rolfe, 69
ionodon Reichb.f., 68
macranthum Lindl., 71
var. hastiferum (Reichbf.
& Warsc.) C.Schweinf.,
70
megalous Schltr., 69
monachicum Reichb.f., 69
obryzatum Reichb f. & Warsce.
71
Pavoni Reichb.f. ex Lindl.,
68
pyramidale Lindl., 71,72
superbiens Reichb f., 72,73
undulatum (HBK.) Lindl.,
738,74
undulatum Salisb., 73
undulatum Sims., 73
undulatum Warner & Williams
12
ventilabrum Reichb, fi &
Warsc., 73,74
Weberbauerianum Krinzl., 64,
65
06-le-da, 89,90,91
OPUNTIA
spp., 159,173,175,179
ORNITHOCEPHALUS
gladiatus Hook., 77
var. peruvianus C. Schweinf.
nr
if
6-s6-pee-ki, 123
OXYTROPIS
sp., 173,179
Patmark, 110
pa-ma, 27
pan, 181,182,196,197,198,199,
203,205,206
pan-chewing, 181,183,184,198,
202
pan dena, 181
pan trinity, 181,198
pangra, 211
pannerale, 212
PARASCHEELIA
anchistropetala Dugand, 112
PERICOME
caudata A.Gray, 174,179
PERSEA
americana Mill., 91
PETALOCENTRUM, 75
angustifolium Schltr,, 76
bicornutum Schltr., 76
pusillum Schltr., 76
PHASEOLUS
vulgaris L., 163,164,177,178
PHYLLANTHUS
fluitans Benth. ex Muell.-Arg.,
29
PIMPINELLA
Anisum L., 207
Pinaceak, 165
pinang, 206
pinang asin, 194
pinang blah, 194
[ xxv ]
pinang kossi, 194
pinang salai, 194
PINUS, 12
edulis Engelm., 159,165,175,
178
ponderosa Dougl. ex P. Laws.,
159,165,178
sp., 165,178
PIPER
Betle L., 188,206
PISTIA
Stratiotes L., 22
platanillo, 116,117
platanote, 44
PLEUROTHALLIS
Broadwayi Ames, 51
gongioglossa Schltr., 52
guadalupensis Cogn., 51
leucantha Schltr., 52
nana A. & S., 51
Sanchoi Ames, 52
Williamsti Ames, 51
POA
Fendleriana (Steud. ) Vasey,
167,178
secunda Presl, 167,178
POPULUS
tremuloides Michx., 170,175,
179
POUTERIA
Ucuqui Pires & Schultes, 87,
90,91
pratie, 285
PROTIUM
sp., 211
PSIDIUM
Guajava L., 212
PTELEA
sp., 178,179
PTEROCARPUS
santalinus L., 193,210
puch-pee-d, 89,91
pudi-adike, 199
puh-pia, 89,90
PUNICA
Granatum L., 212
QUARARIBEA
bracteolosa (Ducke) Cuatr.,32
putumayensis Cuatr., 30
Schultesii Cuatr., 31
QUERCUS, 160
Gambellii Nutt., 170,179
grisea Liebm., 170,175,179
sp., 170,171,179
RaApATEACEAR, 112
RAPATEA
longipes Spruce ex Kérn., 112
115
modesta Maguire, 112,115
Spruceana Kérn, 116
RHAMNACEAE, 160,173
RHAMNUS
sp., 173,179
RHAPHIDORHYNCHUS
Pobeguinii Finet, 257
RHUS
trilobata Nutt. ex T. & G.,
158
RHYTIDANTHERA, 32
magnifica (Gleason) Dwyer,
32,34,35
mellifera R. E.Schultes, 33,34
35
[ xxvii |
splendida (Planch.) van
Tieghm., 34
suleata van Tieghm., 34
RIBES
? aureum, 172,179
sp., 158,172,179
RONDELETIA
rupicola R.Schum., 46
var. chiribiquetana FR. E.
Schultes, 46
Rosacear, 160,172,179
RuBIACEAE, 46
ruktochandan, 210
RUTA
graveolens L., 213
Rutaceak, 160,173
SACCHARUM
oficinarum L., 209,210
SACCOLABIUM
parviflorum Cordemoy, 255
occidentale Krinzl., 254
salei, 211
SALICACEAR, 170
SALSOLA
pestifer 4. Nels., 158
sampig, 212
saora, 210
SAPINDACEAR, 30
satapushpa, 208
saunf, 208
SAUVAGESIA
erecta L., 36
SAXIFRAGACEAR, 172
SCIADOTENIA
toxifera Krukoff & Smith, 120
SCIRPUS
Olneyi A.Gray, 169,175,177,
179
validus Vahl, 169,175,178,
179
sp., 169,179
see-a-se-sep, 42
SENECIO
longilobus Benth., 158
SEQUOIA, 12
SERJANIA
dasyclados Radl., 30
var. sibundoya R. E. Schultes
30
SESAMUM
indicum L., 210
SESBANIA
aculeata Pers., 210
aegyptica Pers., 210
grandiflora Pers., 210
spp., 189
shah-zerah, 208
Shunti, 212
SIGMATOSTALIX, 74
bicornuta Rolfe, 76,77
graminea Poepp. & Endl., 75
hymenantha Sedltr., 75
macrobulbon Krénezl., 75
peruviana Rolfe, 76,77
pusilla Schltr., 76
SIPHONIA
brasiliensis HBK., 79,80,82,
83,85
brasiliensis Willd., 82,83,85
Jjaneirensis (Muell.-Arg. )
Cook, 86
Kunthiana Baill., 79,85
Ridleyana Cook, 81,86
[ xxvii ]
SITANION
Hystrix (Nutt.) J.G.Smith,
167,178
SMILAX
calophylla Wall, 203,208
SOLANACEAR, 44,173
SOLANUM
quitoense Lam., 44,45
sonp, 208
SORGHASTRUM
nutans (L.) Nash, 168,179
SORIDIUM
Spruceanum Miers, 110
SPECKLINIA
graminea Poepp. & Endl., 75
SPHYRARHYNCHUS, 233
SPIRODELA
polyrrhiza (L.) Schleid., 23
SPOROBOLUS
airoides (Torr.) Torr., 158,
168,175,178
Wrightii Munro ex Scribn.,
168,178
STERCULIACEAR, 120
STRYCHNOS
amazonica Krukoff, 41
Erichsonii Rich. Schomb., 41
guianensis (Aub/.) Mart., 42
javariensis Krukoff, 42
Jobertiana Baill., 42
Peckii B.L. Robinson, 43
subcordata Spruce ex Benth.,
43
SUAEDA
suffrutescens S. Wats., 171,
179
su-he-sé-pa, 43
sunnadu-dabbi, 198
supari, 206
su-sé-pe, 43
SWARTZIA
bracteosa Mart. ex Benth., 28
SYZYGIUM
aromaticum (L.) Merr. &
Perry, 206
Cumini (L.) Skeels, 193,209
Jambos (L.) Alston, 212
TAMARINDUS
indica L., 208
tambul, 189,206
tambula kodi, 181,189
tata, 285
tengu, 207
TERNSTROEMIACEAR, 138
THEOBROMA
Cacao L., 133
purpureum Pitt., 133
til, 210
TITHYMALUS
penicillatus Millsp., 2
tobacco, 196,197,203
tombacu, 196,197,202,209
toronja, 45
TRICERATORHYNCHUS,
232
viridiflorus Summerh. , 233,234
TRIDACTYLE
anthomaniaca (Reichb,f. )
Summerh., 236
Scottellii (Rendle) Schltr. , 236
TRIPSACUM, 279
XXVI111
[ ]
TRISETUM
Wolfii Vasey, 167,178
TRIURIDACEAE, 110
TROCHODENDRON, 12
tsau-hess, 111
TYPHA
latifolia L., 166, 169,175,177,
178
TYPHACEAE, 166
tzu-se-hé-pa, 42
ucuqui, 87,89,90,91,92
ucuquirana, 91
UNCARIA
Gambir (Hunt.) Roxb., 196,
209
unde-adike, 193
upakunchica, 206
u-su-se-é-pa, 43
VANILLA
africana Lindl., 221
imperialis Krdnzl., 219,221
polylepis Summerh., 219
ramosa Rolfe, 221
VELLOZIA
phantasmagoria R. Fk.
Schultes, 23
VELLOZIACEAE, 23
VERBENA
ciliata Benth., 158
VERBESINA
sp., 174,179
VIBURNUM
laevigatum, 103
sp., 101
VOCHYSIA
lomatophylla Standl,, 28
obscura Warm., 28
vismiifolia Spruce ex Warm.,
29
VoOcHYSIACEAR, 28
yaupon, 97
YUCCA
baccata Torr., 169,179
spp., 161,169,179
yuct, 91
ZEA
Mays L., 178
ZINGIBER
officinale Rosc., 212
sira, 208
[ xxix |
ERRATA
page 25, line 31
for ramanentibus, read remanentibus
page 115, line 33
for amaz6nica, read amaz6nico
page 134, line 30
replace the comma with a period
page 150, lines 7, 10 and 15
for Matthews, read Mathews
page 195, line 31
for Areca, read Acacia
page 242, line 23
for Kraenzlin, read Kraenzlin
page 249, lines 5 and 9
for E., read A.
page 252, line 32
for H. Perrier, read Perrier de la Bathie
page 253, line 32
for Excell, read Exell
page 281, line 7
for Coons, read Coon
Plate XXXI
for 4, read 5; for 5, read 6; for 6, read 7; add 4 to figure at the
left of 3
Issued April 28, 1951
oe at
BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY
VoL. 14, No. 1
CamsripGr, Massacnusetts, JuNE 27, 1949
DEGRADATION OF PLANT REMAINS
IN ORGANIC SEDIMENTS
BY
Eso S. BARGHOORN
One of the least understood but at the same time fun-
damental processes which operates in nature is the al-
teration of plant residues into economically important
substances such as peat, lignite and coal. In its broadest
terms this degradation of plant substance must be re-
garded as a phase of the major organic cycle of carbon.
It is obvious, however, that in ‘‘geologically permanent”’
large-scale accumulations of organic complexes such as
the fossil fuels, we are concerned essentially with major
deviations from, rather than participation in, the carbon
cycle.
In a certain sense, therefore, the study of the aceumu-
lation of plant residues and their subsequent alteration
comprise an area of botanical investigation which lies
between microbiology on the one hand and the geologi-
cal and chemical aspects of sedimentation on the other.
Microbiological studies of degradation, however, are
more often centered on specific organisms or on the meta-
bolic products of their activity, rather than on an analy-
sis of the effects produced on various naturally occurring
substrata. In addition, emphasis in microbiological stud-
les is quite logically placed more on the degradative as-
pects than on the accumulative aspects of the organic
cycle.
[1]
The ultimate fate of organic residues in nature is de-
termined in large part by diverse factors of the environ-
ment: physical, chemical and biological. The vast bulk
of plant and animal remains is quite rapidly reduced,
through complex energy-releasing processes, to simpler
organic and inorganic compounds; and_ conversely,
through complex energy-consuming processes these are
re-incorporated into the tissues of living organisms. Con-
spicuously active and effective in the biochemical decom-
position changes are the fungi and bacteria.
The rate of degradation due to microbiological activity
is ordinarily most rapid in a warm, moist, highly aerobic
environment. Favorable conditions are probably fulfilled
to the greatest degree on or just below the surface of the
soil. The ratio of accumulation versus degradation in the
soil environment provides variable amounts of the so-
called humus or organic component of the soil. If the
environment is excessively wet, and more particularly
if the availability of oxygen becomes deficient, the rate
of degradation of plant tissues is greatly reduced. Such
conditions reach their extreme in stagnant, poorly
drained and relatively shallow basins such as swamps
and bogs. Under such circumstances the accumulative
phase exceeds the degradative phase and there results a
gradual accretion of modified plant residues. In discuss-
ing the degradation and preservation of plant tissues it
is therefore essential that reasonable distinction be drawn
between environments in which aerobic microbiological
changes may proceed at a rapid rate, as in soil, and sub-
aqueous environments of submergence and oxygen de-
ficiency. This distinction seems to be of fundamental
importance since the character of the degradative changes
is greatly influenced, if not primarily determined, by the
degree of these contrasting conditions of the biological
and physical environment. It might almost be said that
[2]
at one extreme of the spectrum of organic decay there
are the various processes which operate in the soil and
lead to the formation of soil organic matter, while at the
other extreme there are the conditions of anaerobic sub-
mergence. In all cases microbiological processes induce
physical and chemical changes in the plant tissues in-
corporated in the substratum, but the degree of change
and the duration of the change vary profoundly.
Some understanding of the degradation of plant tissue
is essential to a satisfactory interpretation of many types
of plant fossils, since the variously modified remains of
plant parts constitute the bulk of our knowledge of the
paleontological record of plant life. For the most part
these source materials are fragments or occasionally en-
tire plants which fortuitously entered basins of deposition
in which degradative processes were retarded and event-
ually inhibited. Under certain unusual and poorly known
conditions the infiltration of mineral salts in solution and
their subsequent precipitation resulted in an unusually
perfect preservation of the original structure of the plant
tissues. This is well known in the case of many fossil
woods and other plant parts which are silicified or calci-
fied. The sequence of events preceding mineralization,
however, appears to have been basically similar to that
which prevails in any accumulation of plant fragments,
as in the formation of peat and other carbonaceous sedi-
ments. There seems to be no evidence that special envi-
ronmental conditions have been responsible for the ‘‘bio-
logical fixation’’ of mineralized plant fragments, i.e.,
their unusual retention of organic structure. Rather, the
unusual conditions have been those leading to precipita-
tion of the mineral matrix itself, as in the calcareous
nodules, or ‘‘coal balls,’’ of certain coal seams, and the
formation of silicified nodules.
In anticipation of the evidence to be shown later in
[3 ]
this paper, it may be stated here by way of a generali-
zation that: (1) The anaerobic degradation of the plant
cell wall in sediments involves fundamentally similar struc-
tural changes in all plant tissues; and (2) These degrada-
tive changes are directly related to basic physical, chemical
and structural features of the cell wall.
The significant features in the general pattern of struc-
tural change in the anaerobic degradation of wood were
described in a previous study of numerous woods recov-
ered from an archeological site in Boston, Massachusetts
(Bailey and Barghoorn, 1942). Since the publication of
this earlier study, a far wider range of material from
various geological horizons has been examined anatomi-
cally and, in part, microchemically. A more detailed de-
scription of these studies is in the course of publication
(Barghoorn 1949),' but it seems desirable to review here
the salient features of these investigations and their re-
lation to various paleobotanical and anatomical problems
as well as their possible bearing on dating archeological
remains.
In wood of varying age, entombed in mineral or or-
ganic sediments, and permanently submerged, there oc-
curs a gradual reduction of the amount of cellulose con-
tained in the original unmodified cell walls. Depending
on the availability of oxygen during degradative changes,
and its effect on microbiological activity, a variable de-
gree of ‘‘humification’’ results. Stumps of trees, sub-
merged by sudden inundation, commonly occur embed-
ded in peat deposits, yet may show little evidence of the
‘‘humification’’ so characteristic of the peat. For exam-
ple, in the fresh water peats and ‘‘buried forests”’’ of the
‘As a section of the second monograph on the archeology, geology,
stratigraphy and paleobotany of the Boylston Street Fishweir (Papers
of the Robert S. Peabody Foundation for Archeology).
[ 4]
New England coast, now under tidal influence, numerous
stumps and roots may be recovered which retain to a
great extent the color of the original wood, yet whose
tissues are altered to a soft, almost cheese-like consist-
ency. Similarly, the permanently submerged basal parts
of wooden piles driven into sediments in both marine and
fresh water environments show a peripheral zone of de-
graded wood which may be relatively unchanged in color
or gross appearance. Archeological remains of wood pre-
served in the anaerobic environment of estuarine sedi-
ments have been shown to undergo almost no ‘‘humifi-
cation’’ or pronounced change from their original color
(Bailey and Barghoorn, 1942). On the other hand, plant
fragments from autochthonous peats or other organic
accumulations in which aerobic degradation has played
an important part almost invariably exhibit pronounced
changes in color and the other physical and chemical
modifications which feature the concept of ‘‘humifica-
tion’’ (Waksman, 1938).
However, all degraded plant tissues, whether ‘‘humi-
fied” or visibly unaltered, are characterized by a signifi-
cant reduction of their original cellulose, the extent of
loss of cellulose being fundamentally an index of the de-
gree of chemical and physical degradation. In view of
these facts, which are well supported by numerous ana-
tomical and chemical investigations (Mitchell and Ritter,
1934; Jurasky, 1988; Cartwright and Findlay, 19438;
Waksman, 1988; Jahn and Harlow, 1942; etc.), it is of
interest to examine in detail the structural changes which
occur during degradation of the cell wall.
In order to orient the significance of these anatomical
changes it is desirable to note briefly certain fundamental
aspects of the structure of plant cell walls. Among higher
plants the presence of a cell wall is the most conspicuous
visible feature in the organization of tissues, organs and,
[5]
in general, of the entire plant body. The cell wall is com-
posed initially, in its development from pre-existing cells,
of a primary wall, This primary wall is a cellulosic layer
(or series of layers) which ordinarily increases in surface
area and may vary in thickness during the growth and
enlargement of the cell. After growth and enlargement
have ceased, a secondary wall may be formed, always in-
ternal to the primary wall. In many plant tissues, partic-
ularly in reproductive organs, secondary walls are not
formed, and, in such cases the primary wall may become
thick and conspicuous, comprising the bulk of the cell
wall. In the cell walls of wood and woody plant tissues,
however, thick secondary walls are ordinarily developed,
the major topographic and structural feature of such tis-
sues being their greatly thickened secondary walls.
The secondary walls of cells in woody tissues consist,
except in certain cases, of three morphologically distinct
layers or lamellae (Bailey and Kerr, 1985; Bailey, 1988;
1940). The innermost and outermost of these lamellae
comprise relatively thin aggregations of cellulosic wall
substance in which the crystalline cellulose aggregates
tend to be oriented more or less transversely, or in helices
of low pitch. The central and ordinarily thickest layer
of the secondary wall, on the other hand, is composed
most commonly of crystalline cellulose which tends to be
oriented more nearly vertically, or in helices of steep pitch
(Bailey, 1940). All three cellulosic lamellae may be in-
filtrated to varying degrees with encrusting substances
such as lignin, terpenes, resins or other ‘‘protective”’
chemical complexes which retard biological and chemical
degradation.
In the anaerobic degradation of wood it has uniformly
been found that a consistent sequence of degradative
changes occurs. This sequence shows a significant and
fundamental relation to the lamellar structure of the cell
[ 6 ]
wall. The incidence of degradation of the lamellae is in
the following order:
(1) Central layer of the secondary wall
(2) Innermost layer of the secondary wall
(8) Outermost layer of the secondary wall
(4) Primary wall’
Progressive stages in the deterioration of the cell wall
are shown in Plates I-IV. Three outstanding features
may be noted in these thin sections of degraded wood:
(1) areduction of the major portion of the secondary wall
to a granular, virtually amorphous residue; (2) the reten-
tion of a structurally intact cell wall ‘‘layer’’ or ‘‘mem-
brane’’ which corresponds in position to the primary
wall; and (8) the presence of isolated or contiguous
groups of cells whose secondary walls are relatively un-
altered.
That the conspicuous granular remnants of degraded
wall layers are essentially lignin residues may be demon-
strated by their total extraction during delignification
(compare Plate I, Figs. 1, 2 and 8 with Plate III, Figs.
1, 2 and 38). The visibly intact, persistent layers, or oc-
casionally entire cell walls, are truly cellulosic, as may be
shown by their brilliant birefringence in the microscope
when viewed between crossed Nichols. Birefringence in
the thinner cellulosic residual layers is greatly accentu-
ated by delignification.
Because of their extremely tenuous character, certain
structural residues, such as those in Plate I, Figs. 1, 3 and
4 and Plate III, Figs. 1, 2 and 8, might be assumed to
consist solely of primary walls. The composition of these
tenuous ‘‘membranes,’’ however, can be shown to com-
‘The term primary wall refers here specifically to the wall layers
of individual contiguous cells exclusive of the true intercellular sub-
stance which is often erroneously, and confusingly, included as a part
of the ““primary wall’’ (Kerr and Bailey, 1934).
[7]
prise five layers viz., the outermost secondary wall layer
and the primary wall of the two contiguous cells plus the
true intercellular substance between the adjacent primary
walls. The five-layered structure of these tenuous ‘‘mem-
branes”’ is revealed by their relation to the pit border of
bordered pit-pairs. This relationship is most easily ob-
served in the late wood tracheids of coniferous woods, as
shown in Plate III, Figs. 1, 2 and 3 (also compare Plate
ILI, Fig. 8 with Fig. 5). The pit border is solely a struc-
tural feature of the secondary wall. It is apparent, there-
fore, that the tenuous innermost layer of the pit border,
shown in Plate III, Figs. 2 and 8, is the resistant outer-
most or first formed layer of the secondary wall. Hence,
it may be deduced that the persistent fraction of the cell
wall in these tissues consists in large part of the remaining
outermost layers of the secondary wall of two adjacent
cells, including of course the retained and still more ten-
uous primary walls and true intercellular substance. That
the persisting or visibly intact cell wall is actually a very
minor part of the original unmodified secondary wall is
shown in thin sections in which cells possessing intact
secondary walls are intermingled with cells exhibiting
extensively degraded secondary walls (Plate I, Figs. 1
and 2 and Plate III, Fig. 1). The reason or reasons for
the retention of the entire secondary wall in certain cells
which are in direct contact with others in which these
wall layers are extensively degraded remain obscure.
It should be emphasized in connection with its greater
resistance to degradation that the outermost or first
formed layer(s) of the secondary wall possesses quite dif-
ferent staining reactions, optical properties and physical
behavior from those of the broad central layer. The first
formed layer is more heavily lignified than the central
layer and shows far less tendency to swell in strong min-
eral acids, such as 72 per cent sulfuric acid or 40 per cent
[ 8]
hydrochloric acid. It is probable, therefore, that signi-
ficant chemical as well as physical differences exist be-
tween the first formed and the subsequently formed
lamellae of the secondary wall. In these respects the
outermost layer of the secondary wall more closely re-
sembles the primary wall than it does the later formed
lamellae of the secondary wall.
The existence of a pronounced difference in the rate
of degradation of the various cellulosic layers of a single
cell wall presents a seeming paradox in the interpretation
of cellulose decomposition. wo questions immediately
arise: (1) what are the factors, biological or chemical,
which cause the decomposition; and (2) what substance
or substances impart such differential resistance to de-
gradation within the cell wall itself? The fact that dif-
ferential degradation of wall layers is not an isolated
phenomenon, but instead a fundamental feature in the
anaerobic decomposition of plant tissues emphasizes the
significance of these questions and their bearing on the
problem of cellulose decomposition in general.
In a previous study by the author these questions
have been discussed in some detail, particularly with
reference to environmental conditions and the possible
causes of degradation (Barghoorn, 1949). It has been
proposed, though not proven, that the degradation of
wood deeply submerged in marine sediments is probably
due in large part to the hydrolytic breakdown of cellu-
lose rather than directly to microbiological attack.’ It is
probable, although again not proven, that the relatively
high concentration of hydrogen sulfide (and associated
hydro-sulfuric acid) in many carbonaceous marine sedi-
ments is responsible in part for the gradual chemical
'This conclusion does not question the apparently well established
evidence confirming the existence of anaerobic bacteria at considera-
ble depths in peat deposits (Thiessen and Strickler, 1934; Waksman,
1930).
Lo]
hydrolysis of cellulose. Whether similar or comparable
environmental factors are operative in terrestrial carbon-
aceous sediments remains to be determined.
The general uniformity of degradation in wood after
prolonged submergence in sediments of diverse compo-
sition attests to the operation of a widely diffused set of
factors which result in the gradual hydrolysis of the less
resistant cellulosic fractions. In the case of the archeolo-
gical remains previously noted it was found that wooden
stakes, driven through successive strata of marine silt,
peat and glacial blue clay, were uniformly degraded
throughout, the physical condition of the stakes being
identical in their entire length regardless of the surround-
ing media and the duration of submergence (Bailey and
Barghoorn, 1942). Hence, the hydrolytic degradation of
the cellulose appears to have progressed at an approxi-
mately similar rate regardless of the matrix. Similarly,
entire stumps, logs or branches which have been sub-
merged for periods exceeding thousands of years may be-
come uniformly degraded and softened throughout, ex-
hibiting no significant differences between their periphe-
ral and interior parts. Occasionally the innermost por-
tions of larger stumps may retain a core of intact or
incompletely degraded wood. When freshly removed
from its matrix the wood often shows little or no com-
pression failure, nor even any significant change from its
original volume; upon drying, however, it contracts ex-
cessively. Anatomical and chemical study shows that
such degraded wood consists primarily of the lignin res-
idue of the original wood substance, its cellulose content
being reduced to a small fraction of the original, fre-
quently on the order of three to five per cent. The per-
sisting fraction of the cellulose, significantly, however,
is found restricted primarily to those more resistant layers
of the cell wall previously described. All of this evidence
[ 10 ]
indicates that gradual hydrolysis of the cellulosic matrix
of the cell walls has proceeded at a rather uniform rate,
the degradative changes first affecting the peripheral
zone and slowly extending inward.
Ultimate complete degradation of the cellulosic frame-
work of the wood cell wall may be delayed for enor-
mously longer periods than have elapsed since the be-
ginning of post-glacial time (Mitchell and Ritter, 1934;
Jurasky, 1988; Barghoorn and Bailey, 1938). Recent
extensive studies of lignites of Tertiary age show that
in various genera of hardwood trees the lignitized wood
retains from two and one half to six per cent of its origi-
nal carbohydrate fraction, the major constituent of this
fraction being degraded cellulose. In the same deposit
lignitized fruits and seeds may show retention of as much
as 45 to 50 per cent of their original carbohydrate frac-
tion, predominantly cellulose.’
One aspect of the general problem which is of consid-
erable theoretical, if not practical, interest is that dealing
with the causes of selective retention of cellulose in cer-
tain lamellae of the cell wall. Two possible explanations
for the selective degradation of cellulose are immediately
apparent: (1) the presence in the cellulosic matrix of
substances which effectively retard the hydrolysis (either
microbiological or chemical) of cellulose; and (2) actual
chemical differences in the cellulosic framework of the
more resistant lamellae of the cell wall.
The first of these possible explanations finds much sup-
port from various observations and lines of evidence. For
example, it has been established quantitatively by sev-
eral investigators that the degree of lignification corre-
lates with resistance to decay in wood and other woody
tissues (Waksman and Cordon, 1986; Olson, Peterson
' Unpublished data from analyses of the Brandon lignite of Brandon,
Vermont.
[ 11 ]
and Sherard, 1937; Virtanen, Koistinen and Kiuru,
1988). Additional evidence confirming the ‘‘protective’’
effect of lignin has been obtained by the author in a series
of experiments utilizing a species of the cellulolytic bac-
terium Cytophaga. In a series of liquid cultures each
containing mineral salts, filter paper and thin sections of
various woods (Pinus, Sequoia and the hardwood T'rocho-
dendron), it was found that the filter paper was com-
pletely broken down within a period of a week, whereas
the various wood sections were unaffected after six weeks
incubation.
In addition to lignin, other normal constituents of the
cell wall may serve to inhibit biological degradation of
cellulose, viz., resins, terpenes and tanniniferous sub-
stances. Differential resistance of cell wall layers might
logically be ascribed, therefore, solely to the effect of
these encrusting ‘‘protective’’ substances of which lignin
is undoubtedly the most significant in anaerobic degra-
dation of woody tissues. In further support of this ex-
planation of the selective breakdown of the cell wall, it
should be emphasized that the most resistant layer of the
secondary wall, viz., the outermost or first formed layer,
is often far more heavily lignified than the broad central
layer (Bailey and Kerr, 1985). Moreover, the primary
wall, the most resistant of all the lamellae of the cell
wall, is intensely lignified (Ritter, 1925; Kerr and
Bailey, 19384).
The effect of degrees of lignification in different lamel-
lae of the cell wall often produces striking morphological
effects on the mode of attack of wood by various fungi.
Fungi belonging to diverse genera of the Pyrenomycetes
and Fungi Imperfecti commonly degrade wood by dis-
solving the central and often the innermost layers of the
secondary wall, but leaving the outermost layer and the
primary wall visibly unaltered (Bailey and Vestal, 1937;
[ 12 ]
Barghoorn, 1944). Early stages of this restricted break-
down of the cellulosic matrix are illustrated in Plate IV,
Figs. 1 and 2, showing the invasion of wood by fungal
hyphae. Similar in nature are the effects of certain brown
rots on the degradation of wood. These are shown, per-
haps more diagrammatically in Plate IV, Figs. 8 to 6.
In some cases the entire secondary wall (with the excep-
tion of the tenuous outer layer) is completely removed,
leaving a structural residue consisting of five layered
‘‘membranes’’ similar in composition to those previously
described from wood degraded under anaerobic condi-
tions (compare Plate III, Figs. 1 and 8 with Plate IV,
Figs. 3, 4, 5 and 6).
All of this evidence indicates that the deterioration of
wood is primarily a process of removal of cellulose, the
rate of loss being, in many cases, significantly correlated
with the degree of lignification or the protection of cel-
lulose by other substances.
A fundamental exception to this relatively simple ex-
planation, however, may be determined by an anatomical
study of plant remains extracted from peats. In defloc-
culated, and subsequently delignified (‘‘dehumified’’),
samples of fibrous peats innumerable delicate plant frag-
ments are often released. In grass and sedge peats these
fragments are most commonly cellulosic epidermal resi-
dues of roots and rhizomes. Microscopic examination
often reveals preservation of even the most delicate epi-
dermal structures such as root hairs, root cap cells and
epidermal papillae (Plate VI, Figs. 1 to 6). Although
these tenuous residues of roots have retained minute ana-
tomical features in certain parts, they are devoid of their
thick walled fibrous or conductive tissues (Plate VI,
Figs. 1 and 2). However, in undelignified preparations
of comparable roots there may be found degraded rem-
nants of the originally thick walled conducting tissues
[ 13 ]
and fibers, whose secondary walls have been reduced to
granular residues. The secondary wall residues and other
degraded or ‘‘humified’’ decomposition products are
readily extracted (with little or no deleterious action on
the remaining cellulose) by means of various techniques
utilizing sodium chlorite as the delignifying agent
(Barghoorn, 1948).
Studies of fibrous peats of varying post-glacial age
consistently show a cellulosic residue, often representing
avery small fraction of the original sample. The amount
of cellulose recovered is much influenced by the tech-
nique employed in lignin extraction; direct repeated
chlorination or treatment with acidified hypochlorite so-
lutions may result in oxidation and partial (or complete)
solution of the persisting degraded cellulose residues.
Anatomically, however, structural cellulosic residues pre-
served in peats consist for the most part of thick, previously
unlignified primary cell walls. Tissues possessing thick,
lignified secondary cell walls may undergo extensive de-
gradation in environments in which even delicate cellu-
losic cell walls are incompletely broken down. In recog-
nition of these facts it seems a paradox of note that the
most delicate tissues of various plant organs may be far
more resistant to anaerobic degradation than are cells or
tissues possessing thick, frequently heavily lignified sec-
ondary cell walls.
In view of these observations, which are consistently
supported by anatomical studies as well as by chemical
analyses of degraded plant remains, it seems quite diffi-
cult to interpret selective degradation of different por-
tions of the plant cell wall except in terms of chemical
differences in the successively formed lamellae of the cell
wall. In other words, the ewceptional resistance of the
primary cell wall to degradation may be due to chemical
rather than physical ‘‘protective’’ factors. Such an inter-
[ 14 ]
pretation is strongly supported by the fact that primary
wall cellulose often persists after degradation of the cel-
lulose of heavily lignified layers of the secondary wall.
For this and other reasons it seems difficult to interpret
selective degradation of cellulose in different parts of the
plant cell wall except in terms of chemical differences and
resistance to hydrolysis in successively formed lamellae.
The persistence of the primary and occasionally the
outermost layers of the secondary cell wall is strikingly
shown in many silicified woods. Not uncommonly, sili-
cified wood, regardless of geologic age, contains unusual
amounts of organic residues, occasionally present to such
an extent that the demineralized wood may be embed-
ded, sectioned and stained much as living tissue (Arnold,
1981; 1941). In such material of diverse groups of
plants which may range in age from Devonian to Ter-
tiary, there is no cellulose remaining; it seems likely that
silicification took place at a time when degradation of
the cell wall had not yet passed the stage of the partial
retention of a cellulosic structural residue. In other
words, the cellulosic framework of the tissues was re-
tained for a sufficient length of time to allow preservation
of structure before silicification began. After or during
the silicification process the remaining cellulose was lost
from the tissue, leaving a modified but coherent lignin
residue. ‘This modified and silicified residue simulates
the original cellulosic residue of anaerobically degraded
wood (compare Plate I, Figs. 1 and 8 and Plate II, Figs.
2 and 8 with Plate V, Figs. 1, 2 and 5).
Whether the greater resistance to degradation of the
cellulosic lamellae of the primary wall and the outermost
secondary wall of plant cells is due to intrinsic chemical
factors or primarily to physical protection by extraneous
non-cellulosic substances cannot be determined without
further codrdinated histological and chemical study.
[15 ]
However, it is evident from anatomical and microchem-
ical investigation that resistant fractions of cellulose oc-
cur in many organic deposits of various geologic ages and
that these resistant fractions are directly related to pre-
determined structural features of the plant cell wall;
hence they have their basis in the biochemistry of growth
and differentiation of the cell wall.
Since the ultimate loss of cellulose in plant remains
may, under certain conditions, be delayed to varying
degrees for immense periods of geologic time, it is appar-
ent that simple generalizations can scarcely be made re-
garding the complex processes of degradation in organic
sediments as a whole. Chemical analyses of peat and
humus have supplied data showing gross trends in the
chemical alteration of plant remains in deposits (W aks-
man, 1938). However, such empirical analyses do not
correlate microchemical and anatomical aspects of degra-
dation, and in general they interpret natural decompo-
sition of plant remains in terms of plant substance rather
than in terms of the complex organization of cells and
tissues. It is significant, however, that the less favorable
the conditions are for microbiological processes, the
greater is the retention of the original cellulose in plant
accumulations. With increasing geologic age, total loss
of cellulose occurs through ‘‘coalification’’ and various
changes associated with the transformation of plant resi-
dues into higher rank coals. The problems of the earlier
stages of cellulose degradation in anaerobic environments
and of the processes of ‘‘humification’’ in general are
in need of further co6rdinated anatomical, microbiologi-
cal and chemical studies. Such investigations will be es-
sential before a clear understanding of the major biologi-
cal and chemical changes occurring between plant source
materials and their geologic accumulation in the form of
fossil fuels will be possible.
[ 16 ]
me
SUMMARY
. Alteration of plant residues under anaerobic condi-
ditions in organic sediments is a fundamental but
inadequately known phase of the accumulation of
carbon in nature.
Details of anatomical changes during decomposition
of plant tissues are of importance in the morpholo-
gical interpretation of many structurally preserved
plant fossils.
. Anaerobic degradation of the plant cell wall involves
certain basic structural changes which appear to be
similar in all plant tissues; these changes are directly
related to basic physical and chemical features in
the organization of the cell wall.
Degradation of the cell wall, either aerobically or
anaerobically, is characterized primarily by loss of
cellulose; the removal of cellulose most frequently
follows a uniform sequence determined by the lamel-
lar organization of the wall.
The incidence of degradation of cell wall lamellae in
lignified tissues occurs in the following sequence:
1. Central layer of the secondary wall
2. Innermost layer of the secondary wall
Outermost layer of the secondary wall
. Primary wall
eo
The greater resistance of certain lamellae of the cell
wall may be logically assigned to protection of the
cellulosic framework by lignin and other ‘‘encrust-
ing’’ substances. The persistence of unlignified pri-
mary walls in fibrous peats, however, cannot be ex-
plained by such mechanical protection, and demands
an alternate explanation.
[17 ]
7. It is proposed, although it cannot be demonstrated
within the scope of this study, that the greater re-
sistance to hydrolysis of the primary wall and the
outer secondary wall layeris due to chemical differ-
ences in the organization of the cellulosic matrix,
rather than solely to a physical relation with lignin
and other protective complexes.
8. The ultimate loss of cellulose in organic deposits
may be delayed for periods measured in terms of
geologic time, as shown by its presence in lignites
of Tertiary and older age. The rate of loss, however,
is greatly influenced by the initial biological condi-
tions of deposition as well as by subsequent environ-
mental changes accompanying ‘‘coalification. ”’
9. Mineralization of plant remains apparently often oc-
curs at a stage when cellulosic structural residues are
still retained. Infiltration and precipitation of min-
erals, such as silica and calcium carbonate, occasion-
ally cause a ‘‘fixation’’ of structure which, in min-
eral form, retains a large measure of biological detail.
Such petrifactions may or may not retain a rather
high percentage of the original, though chemically
modified, organic matter during subsequent geologic
change.
9
10. Many aspects of the problem of ‘‘humification’”’ and
‘*eoalification’’ of plant residues are known only in
empirical terms. An adequate understanding of
these complex processes will necessitate extensive
coordinated microbiological, anatomical and chemi-
cal investigation.
Institute for Research in General Plant Morphology
Harvard University
[ 18 |
LITERATURE CITED
Arnold, C.A. 1931. On Callixylon Newberryi (Dawson) Elkins et
Wieland. Contrib. Museum of Paleontology, Univ. of Mich. 3:
207-232.
—— 1941. The petrifaction of wood. The Mineralogist. 9: 323-324;
SOO=SI0O%
Bailey, I.W. 1938. Cell wall structure of higher plants. Indust. Eng.
Chem. 30: 40-47.
—— 1940. The walls of plant cells. Amer. Assoc. Adv. Sci. Publ.
no. 14; 31-48.
and E. S. Barghoorn. 1942. Identification and physical condi-
tion of the stakes and wattles from the fishweir. (In Johnson et al. :
The Boylston Street Fishweir.) Papers of the Robert S. Peabody
Foundation for Archeology. 2: 82-89.
and T. Kerr. 1935. The visible structure of the secondary wall
and its significance in physical and chemical investigations of trach-
eary cells and fibers. Journ. Arn. Arboretum 16: 273-300.
and Mary Vestal. 1937. Significance of certain wood-destroying
fungi in the study of the enzymatic hydrolysis of cellulose. Journ.
Arn. Arboretum 18: 196-205.
Barghoorn, E.S. 1948. Sodium chlorite as an aid in paleobotanical
and anatomical study of plant tissues. Science. 107: 480-481.
—— 1949, Paleobotanical studies of the fishweir and associated de-
posits. Papers of the Robert S. Peabody Foundation for Archeology.
(in press)
and I. W. Bailey. 1938. Occurrence of Cedrus in the auriferous
gravels of California. Amer. Journ, Bot. 25: 641-647.
and David Linder. 1944. Marine fungi: Their taxonomy and
biology. Farlowia. 1: 395-467.
Cartwright, K. St. G. and W. P. K. Findlay. 1943. Timber decay.
Biol. Reviews. 18: 145-158.
Freudenberg, K. 1932. The relation of cellulose to lignin in wood.
Journ. Chemical Education. 9: 1171-1180.
Jahn, E.C. and W.M. Harlow. 1942. Chemistry of ancient beech
stakes from the fish weir. (In Johnson et. al.: The Boylston Street
Fishweir) Papers of the Robert S. Peabody Foundation for Arche-
ology. 2: 90-95.
[ 19 ]
Jurasky, K.A. 1938. Wandlungen des Holzes auf dem Weg zur Kohle.
Geologischen Rundschau. 29: 441-461,
Kerr, T. and I. W. Bailey. 1934. The cambium and its derivative
tissues. No. X. Structure, optical properties and chemical compo-
sition of the so-called middle lamella. Journ. Arn. Arboretum. 15:
327-349,
Mitchell, R. L. and G. J. Ritter. 1934. Composition of three fossil
woods mined from the Miocene auriferous gravels of California.
Journ. Amer. Chem. Soc. 56: 1603-1605.
Olson, F. R., W. H. Peterson and E. C. Sherard. 1937. Effect of lig-
nin on fermentation of cellulosic materials. Ind. Eng. Chem. 29:
1026-1029,
Ritter, G.J. 1925. Distribution of lignin in wood, Ind. Eng. Chem.
17: 1194-1208.
Thiessen, R. and H.S. Strickler. 1934, The distribution of micro-
organisms in four peat deposits. Coop. Bull. no. 61. Mining and
Metallurgical Investigations U.S. Bureau of Mines, Pittsburgh,
Pennsylvania.
Vertanen, Koistinen and Kiuru. 1938. Suomen Kemistilehti. 11: 30
(not seen, abstract in Papier Fabrik 38: 22, 1940).
Waksman, Selman A. Humus. 1938. pp. 93-129; pp. 261-288.
Williams and Wilkins, Baltimore.
——— 1930, Chemical composition of peat and the réle of microorgan-
isms in its formation. Amer, Journ. Science, 19: 32-54.
and T. C. Cordon. 1938. On the nature of lignin. Journ. Amer.
Chem. Soc. 58: 969-972.
ILLUSTRATIONS
EXPLANATION OF THE ILLUSTRATIONS
Piare I. Fia. 1. Transverse section of the wood of a sub-
merged white pine stump, age about 2000 years, showing
differential degradation of the cell wall. Note the presence
of some intact cell walls, others incipiently degraded and
many cells in which the secondary wall is largely reduced to
a lignin residue occupying the center of the lumen, 70.
Fic. 2. Transverse section of the same material showing
incipient degradation of the secondary wall of an isolated
tracheid (center) surrounded by cells in which the secondary
wall has been largely degraded. 170.
Fic. 38. Transverse section of the same, but more highly
magnified. Note the discrete mass of the ‘‘coagulated’’ rem-
nants of the major part of the secondary wall in the center
of each cell. 870.
Fic. 4. Tangential longitudinal section of the same material.
The dark, heavily stained amorphous material represents the
degraded remains of the central and inner layers of the sec-
ondary walls. 100,
Fic. 5. Transverse section of a spruce pile, submerged in
marine silt for 100 years. This section from the outer rings
of the pile shows incipient degradation of the central and
inner layers of the secondary wall. 500.
Fic. 6. Transverse section from a different part of the white
pine stump shown in Fig. 1. Note the complete breakdown
of the central and inner layers of the secondary wall and the
retention of the primary wall and the outermost layers of the
secondary wall. 500.
PLATE [
EXPLANATION OF THE ILLUSTRATIONS
Pirate Il. Sections of wood of submerged white pine stump.
Fic. 1. Transverse section. Note the incipient degradation
of the inner lamellae of the secondary wall of the tracheids,
which shows compression wood, or “‘rotholz’” structure. The
heavily stained layers between the degraded inner lamellae
and the less heavily stained outer secondary wall are non-
cellulosic layers of the secondary wall. »* 500.
Kia. 2. Transverse section. The visibly intact layers of the
cell wall consist of the intercellular substance, primary wall
and the outer secondary wall. The lignin residues of the
central and innermost layers of the secondary wall form a
discrete cylinder in the lumen of each cell. In this section,
as also in Fig. 8, the outermost secondary wall is unusually
thick and appears to consist of more than one layer. Unde-
lignified, stained with Ruthenium red. “800,
Kia. 3. Same section shown in Fig. 2, except more highly
magnified, Undelignified, stained with Ruthenium red,
1400,
Kia. 4. Transverse section showing varying extent of de-
gradation of the secondary wall lamellae. ~ 800.
Fic. 5. Transverse section of less degraded wood from inner
parts of the stump. Note the varying degrees of degradation
of the secondary wall and the retention of a structural frame-
work of modified cell walls. Unstained preparation. 220.
kia. 6. Same specimen as shown in Fig. 5, but from more
extensively degraded wood. Unstained preparation, 530,
PLATE
EXPLANATION OF THE ILLUSTRATIONS
Prare IIL, Fic. 1. Transverse section of wood of white pine
stump. The section shows the cellulosic structural residue
of degraded wood after extraction of lignin and lignin resi-
dues by sodium chlorite treatment. Four tracheids possess
visibly intact secondary walls. All other cells are repre-
sented by walls consisting of intercellular substance, primary
walls and outermost secondary walls. Stained with Rutheni-
um red. 500,
Fig, 2. Same specimen as Fig. 1. The cellulosic structural
residue consists of intercellular substance, primary walls and
outermost secondary walls. Evidence for the presence of a
secondary wall residue is provided by the persisting first
formed layers of the pit borders. Delignified section stained
with Ruthenium red. * 870.
Fic. 3. Same specimen as Fig. 1. Evidence that the cellu-
losic residue consists in large part of the outermost layer of
secondary wall is shown by the conspicuous remnants of the
borders of bordered pit-pairs. Note the retention of the pit
torus. Delignified, stained with Ruthenium red. » 870.
Fie. 4. Tangential longitudinal section of a degraded late
wood tracheid of white pine showing the granular lignin
residue of the secondary wall, Note the remnants of the
bordered pit-pairs. Such cell wall residues, when delignified
and viewed in transverse section, show the organization rep-
resented in Figs. 1, 2 and 8. * 500.
Fic. 5. Bordered pit-pairs of a normal spruce tracheid. Com-
pare the lamellae with those shown in Figs. 2 and 3.) 870.
Fic. 6, Transverse section of “‘rotholz’’ of degraded white
pine wood after treatment with 72 per cent sulphuric acid.
The more heavily lignified primary walls and outer secondary
walls yield a coherent structural residue comparable to that
Ys
shown in Plate II, Figs. 2 and 8.» 800.
kia. 7. Same specimen as shown in Fig. 6, but from differ-
ent portion of the stump. Section treated with 72 per cent
sulphuric acid. Note the lignin residue of the pit borders
and compare with the cellulosic residue of the pit borders
shown in Fig. 8.0 500.
ae Be
PLATE
EXPLANATION OF THE [ILLUSTRATIONS
Prare IV. Fie. 1. Transverse section of xylem of Laurelia
aromatica Juss., showing enzymatically produced cavities in
the secondary wall. Note the partial localization of action
within the central layers. (After Bailey, I. W. and Mary
Vestal: Journ, Arnold Arboretum 28, Plate 209: 19387.)
~990,
kia, 2. Transverse section of hard pine wood which was ex-
posed to the sea and to the action of marine fungi. Enzy-
matic attack on the cell wall is concentrated within the cen-
tral layers. Compare with anaerobic degradation as shown
in Plates I, Il and III. 500.
Fic. 8. ‘Tangential section of xylem of U/mus crassifolia
Nutt. showing normal wood structure. * 100.
Kia, 4. Tangential section of xylem of Ulmus americana L.
degraded by an unidentified brown rot fungus. The structu-
ral residue closely resembles delignified residues of anaero-
bically degraded wood as shown by comparison with Plate
III, Figs. 1, 2 and 8. 100.
Fra, 5, ‘Transverse section of xylem of U/mus americana LL.
showing normal wood, with scattered gelatinous fibers.
LOO.
Fic. 6. Transverse section of xylem of Ulmus americana L.
degraded by unidentified brown rot. Same specimen as
shown in Fig. 4. The structural residue is lgnified cellulose
as shown by lignin extraction, ~ 100,
PLatrE IV
Gy
os
EXPLANATION OF THE ILLUSTRATIONS
Prare V. Fie. 1. Transverse section of an unidentified fos-
sil wood of Cretaceous age. The wood is an intensely silici-
fied organic residue of the original woody tissue. The cell
wall layers retained closely resemble the cellulosic residues
of anaerobically degraded wood as shown by comparison with
Plates I and II. Ground section. » 182.
Kia. 2, Same specimen as shown in Fig. 1, but more highly
i y @6y. . ry : .
magnified. Note the “lignin’’ residue of the secondary wall.
732.
Fig. 38. Robinia Pseudo-Acacia LL. Portion of vessel wall
showing structure of the vestured pits. The vestured portion
of the pit consists of primary wall. ~< 1532,
hic. 4. Robinia sp. Silicified wood from the Tertiary of Mon-
tana. Portion of vessel wall showing retention of structure
of the vestured pits. Compare with normal wood of Robinia
Pseudo-Acacia L. shown in Fig. 8. Ground section. 860.
Fie. 5. Same specimen as shown in Figs. | and 2, but with
varying degrees of degradation of the cell wall previous to
silification. Ground section. 732.
Kia. 6. Tangential longitudinal section of same specimen as
shown in Fig. 4. The beaded structure of the more resistant
primary wall is preserved as an organic residue, consisting
of modified lignin and embedded in silica. Ground section.
225,
Fig. 7. Same as Fig. 6, except more highly magnified.
Ground section, » 860,
\
PLATE
EXPLANATION OF THE ILLUSTRATIONS
Prare VI. Fic. 1. Portion of root of Juncus sp. isolated from
postglacial peat. Age approximately 4000 to 5000 years.
The heavily lignified fibrous and conductive tissue in the
center of the root has been largely degraded, whereas the
delicate unlignified epidermal structures are preserved.
132.
ia, 2. Root epidermis of Scirpus sp. from same peat. Note
preservation of epidermal cells. 126.
Fig, 3. Root epidermis of Carex sp. from same peat. * 126.
Fic. 4. Root tip of unidentified monocotyledonous root from
same peat. Note preservation of minute structural details of
root cap. ‘Treated with acidified sodium chlorite. ~ 126,
Fig. 5. Cells of a root apex meristem from the same peat
showing preservation of primary cell walls and the residue
of cell contents. Thin section of peat, unstained, 500.
Fic. 6. Root epidermis of Carex sp. showing preservation of
unlignified primary walls of epidermal cells. 126.
PLATE VI
[4 L
pres bate
ae :
ue
Re eae ae Be OS Bess
Camprinar, Massacnuserts, Ocroper 26, 1949
BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY
PLANTAE COLOMBIANAE XII
DE PLANTIS PRINCIPALITER AMAZONIAE
COLOMBIANAE INVESTIGATIONES
BY
Ricuarp Evans Scuuutres!
THE Amazonia of Colombia, vast and, for the most
part, still unknown, has a surprisingly varied flora. Bo-
tanical explorations of this northwesternmost part of the
great Amazon Valley have been pitifully few in relation
to the size of the area and the wealth of its flora. Colom-
bia’s Amazon region is divided into the Comisarias of the
Putumayo, Caqueté, Amazonas and Vaupés. It com-
prises about 404,000 square kilometers or one-third the
area of the entire country. It is at once obvious how
vital to our understanding of the flora of the rest of Co-
lombia, as well as of the whole Amazon basin, is a clear
conception of the phytogeographical relationships of the
vegetation of this immense area.
In the course of identifying recent collections from the
Colombian Amazonia, I have found that a number of
these represent hitherto apparently undescribed concepts
or concepts previously unknown either from Colombia
or from its Amazonian regions. It seems advisable to
publish, from time to time, collected notes of this type
' Botanist, Bureau of Plant Industry, Soils, and Agricultural Engi-
neering, Agricultural Research Administration, United States Depart-
ment of Agriculture; Research Fellow, Botanical Museum, Harvard
University.
[ 21 |
Voi. 14, No.
as a contribution towards our understanding of the flora
of Colombia.
I wish to thank the following botanists for their collab-
oration: Dr. José Cuatrecasas of the Chicago Museum,
who has studied the representatives of Cecropia and
Quararibea; Dr. Robert E. Woodson of the Missouri
Botanical Garden, who identified the apocynaceous spe-
cies; Padre Lorenzo Uribe of Medellin and Bogota,
Colombia, who determined the specimens of Inga; Dr.
B. A. Krukoff and Mr. Joseph Monachino who have
attended to the collections of Strychnos; and Dr. F. A.
Stafleu of Proefstation voor de Java Suikerindustrie,
Pasoeroean, Java, who identified the representatives of
Vochysia. 1 have also to thank Dr. Frederick J. Hermann
of the United States Department of Agriculture for
placing at my disposal the notes and identified plant
specimens which he collected, some for the first time in
Colombian territory, in the Leticia area in 1944.
CYPERACEAE
Fimbristylis miliacea (L.) Vahl Enum. 2 (1805)
287.
Collections of Mimbristylis miliacea have been made
in British and Dutch Guiana, Venezuela and Ecuador.
Hermann 11318 represents apparently the first collection
to be reported from Amazonian Colombia.
Cotomsra: Intendencia del Amazonas, Leticia, April, 1944, Fred-
erick J. Hermann 11318.
ARACEAE
Pistia Stratiotes Linnaeus Sp. Pl. (1758) 963.
This very widespread tropical American plant is known
from almost all parts of Middle and South America. Al-
though collections have been made from nearly every
other district of Colombia, Hermann 11301 would seem
[ 22
to be the first reported from the Amazon watershed of
the Republic.
CotomsrA: Intendencia del Amazonas, Rio Loretoyacu, floating on
the river near its mouth, April, 1944, Frederick J. Hermann 11301.
LEMNACEAE
Lemna valdiviana Philippi in Linnaea 33 (1864)
239.
The collections cited below seem to be the first of this
minute floating plant from the Amazon watershed of
Colombia and adjacent Peru.
Cotompia: Intendencia del Amazonas, Rio Hamacayacu, April,
1944, Frederick J. Hermann 11312.
Peru: Departamento de Loreto, Rio Amazonas, Isla Cacao, April,
1944, Frederick J. Hermann 11313.
Spirodela polyrrhiza (L.) Schleiden in Linnaea 18
(1839) 392.
This interesting little aquatic plant has apparently
never before been collected in the Colombian Amazon
watershed. It has been known from Boyacé and Santa
Marta in Colombia and from Guayas Province in Ecua-
dor and Loreto in Peru.
Cotompia: Intendencia del Amazonas, Leticia, April, 1944, Fred-
erick J. Hermann 113165.
VELLOZIACEAE
Vellozia phantasmagoria PR. I. Schultes in Bot.
Mus. Leafl. Harvard Univ. 12 (1946) 130, pl. 19, 20.
A specimen of Cuatrecasas 7700 has recently come to
my attention during studies in the United States National
Herbarium. It would appear to represent Vellozia phan-
tasmagoria, a dominant shrub on Mount Chiribiquete
which is located slightly southeast of San José del
Guaviare and which has a number of floral elements in
common with the granitic hill at San José. The sterile
[ 23]
condition of the specimen, however, precludes definite
identification. Nevertheless, it represents the northwest-
ernmost collection of the Velloziaceae from the Amazon
Valley and from South America.
Cotompra: Intendencia del Meta, Rio Guaviare, San José del Gua-
viare, terrenos graniticos, sabana, alt. 270 m. **Ramificada, 1-2 m.
alt.’’ December 11, 1939, J. Cuatrecasas 7700.
MoRACEAE
Cecropia mocoana Cuatrecasas sp. nov.
Arbor parva. Ramuli scabri, pilis conicis rigidis in-
clinatis et pilis longioribus hamatis praediti. Folia ampla,
membranacea, peltata, longe petiolata. Petiolus robus-
tus, 50 em. longus, striatus, copiose hirsutus cum pilis
longis saepe hamatis. Lamina palmata, profunde (8/4—
4/5) 8-lobata; lobis majoribus obovato-oblongis, minori-
bus obovato-subrotundatis, omnibus abrupte acuminatis,
basi contractis, margine integra vel leviter irregulariterque
sinuata, lobo maximo ab insertione petioli usque ad api-
cem 54cm. longo et 30 cm. lato, lobo minimo 24 em.
longo, 15 cm. lato, parte concrescenti 10-19 cm; supra
viridis, strigosa, pilis conicis brevibus et pilis longis acu-
tissimis inclinatis munita, nervis principalibus et secun-
dariis notata; subtus albida, nervis primariis crasso-
eminentibus valde hirsutis, secundariis angulo obtuso
1-3.5 cm. distantibus prominentibus hirsutulis, tertiis
roseis transversalibus et reticulatis demum hirsutulis, in-
ter hos lamina dense albo-arachnoideo-tomentosa. In-
florescentiae masculae geminatae. Pedunculus rigidus,
mediocris, 6.5-7 cm. longus, strigosus, pilis hamatis acu-
tis satis armatus. Spatha ovoideo-oblonga, acutiuscula,
circa 13 cm. longa, extus villoso-hirsuta, intus puberula,
Amenta masculina circa 28 fasciculata, graciliter pedi-
cellata, praefloratione 7-10 cm. longa, postea usque ad
17 em. longa, 3 mm. lata. Perigonium tubulosum, 2
[ 24 ]
mm. longum, glabrum vel apice setis parcissimis minutis
donatum. Pedicelli subcapillares, 12-20 mm. longi,
dense scabro-hirsutuli, pilis hamatis instructi.
Dr. Cuatrecasas writes: ‘*‘Cecropia mocoana is closely
related to C. marginalis Cuatr. and C. discolor Cuatr.
From both, it differs in having leaves which are a little
thinner and more abundantly pubescent on the upper
surface and in having more conspicuous scales (small
conic hairs) which cause the leaves to be rough. The
leaves on the under surface are more like those of C.
marginalis because of the nerves and venation which are
more or less hirtose and not covered with arachnoideous
tomentum; in C. discolor the arachnoideous tomentum
is brightly white and covers the network. The spathes
of Cecropia mocoana are densely villous-hirsute as in C.
discolor (in this species they are glabrous or only puberu-
lent inside, contrary to the statement in the original de-
scription), whereas in C. marginalis they are covered
with a very short and obsolete tomentum. Only further
abundant collections can indicate whether Cecropia mo-
coana represents a male specimen of one of these earlier
species, ”’
Cotoms1a: Comisaria del Putumayo, Mocoa, alrededores al norte,
750-850 m. ‘‘Small tree.’? December 38-7, 1942, Richard Evans
Schultes & C. Earle Smith 2003 (Tyre in Chicago Museum).
Cecropia porvenirensis Cuatrecasas sp. nov.
Arbor magna. Ramuli seabri, pilis conicis et longiori-
bus hamatis satis instructi. Folia membranacea vel sub-
coriacea, peltata. Petiolus 32 em. longus, juvenilis albo-
arachnoideus, deinde hirtus, pilis longis tenuibus rigidis
patentibus inferne hamatis tantum ramanentibus. La-
mina palmata profunde (2/38) 7-vel 6-lobata, lobis obo-
vatis vel obovato-oblongis, basi constrictis, apice abrupte
acuminatis, margine integra vel leviter irregulariterque
[ 25 ]
sinuata, lobo maximo ab insertione petioli usque ad api-
cem 85 cm. longo, 15 em. lato, lobo minimo 14 cm.
longo, 8 cm. lato, parte centrali concrescenti 7-13 cm. ;
supra viridis, scabra, pilis conicis minutis satis atque
tuberculis acutis et pilis sparsis longis praedita, nervis
primariis secundariisque valde conspicuis; subtus alba,
nervis principalibus in foliis juvenilibus tomento arach-
noideo deciduo deinde sparse hirtis, nervis secundariis
prominentibus, adscendentibus, 1-2 cm. distantibus,
plusminusve arachnoideis et sparsissime hirtulis vel gla-
brescentibus, reticulo venularum obsoleto et lamina re-
liqua tomento arachnoideo albo dense tecta. Stipula in
statu juvenili 8 cm. longa, acuta, villoso-hirta, margine
utrinque glabra, intus villoso-sericea. Inflorescentiae
femineae geminatae. Pedunculus 6-8 cm. longus (fructi-
fer usque ad 17 em. longus) scabrido-hirtulus, tuberculis
minutis atque pilis rigidis fragilibus patentibus rectis et
hamatis munitus. Spatha cylindrica, apice longe angus-
tata, acuta, 16 cm. longa, extus sparse puberula et arach-
noidea, leviter scabrida, intus pubescenti-arachnoidea.
Amenta feminea vulgo 5 vel 6 (4-7), leviter pedicellata,
11 cm. longa, 4 mm. lata, fructifera in sicco usque ad
23 cm. longa, 7-8 mm. crassa. Perigonium minute tu-
berculatum. Achaena elliptica, 2 mm. longa, 1 mm.
lata, laevia. Pedicelli crassi, 3-4 mm. longi, scabri, pilis
conicis brevibus et pilis longioribus hamatis instructi.
Concerning the relationship of this new species, Cua-
trecasas writes: ‘‘Cecropia porvenirensis is closely related
to C. marginalis Cuatr., but it differs by having the upper
surface of the leaves very scabridous, the under surface
more densely white-arachnoideous with the nerves much
less hispid, and the lower part of the petiole with hooked
hairs; the amenta are pedicellate, longer and usually five
or six, while in C. marginals they appear in groups of
four and are sessile. The stipule of Cecropia porveniren-
[ 26 |
sis is villose inside, whereas in C. marginalis it is gla-
brous.”’
Cotomsia : Comisaria del Putumayo, Puerto Porvenir cerca de Puerto
Ospina, alt. 250 m. “‘Large tree. Yarumo.’’ May 20, 1942, Richard
Evans Schultes 3697 (Tver in Chicago Museum).
LEGUMINOSAE
Inga macrophylla Humboldt & Bonpland ex
Willdenow Sp. Pl. 4 (1806) 622.
Inga macrophylla, which ranges from Venezuela to
Peru and Bolivia, was first reported from Colombia by
Uribe (in Caldasia 8, no. 14 (1945) 347), on the basis of
material from the Intendencia del Meta. The second and
third Colombian collections, cited below, greatly extend
the known range of this species in Colombia. In the
Colombian Amazonia, this species, cultivated for the
edible pulp around the beans, is called guava machete or
guamo machete. In the Tikuna Indian language, it is
known as pa-md.
Cotompr1a: Comisaria del Amazonas, Rio Karaparand, between E]
Encanto and the mouth, May 22-28, 1942, Richard Evans Schultes
3857.—Comisaria del Amazonas, Leticia. ‘‘Bushy shrub. Flowers
white. Fruit long. Guamo machete. Tikuna: pa-md’’. September 20,
1945, Richard Evans Schultes 6542.
Inga setifera de Candolle Prodr. 2 (1825) 432, 615.
According to Uribe (in a letter to the writer), Inga
setifera has not hitherto been reported from Colombia.
It is known to be common in the Departamento de
Loreto in Peru. In Leticia, the plant is called chimbillo
by the Spanish-speaking population, kau-ré by the
Tikuna Indians.
CotomBia: Comisaria del Amazonas, Leticia. “‘Flowers yellow.
Pulp of fruit edible. Chimbillo. Tikuna: hau-ré.’’ September 20,
1945, Richard Evans Schultes 6543.
[ 27 |
Swartzia bracteosa Martius ew Bentham in Mar-
tius Fl. Bras. 15, pt. 2 (1870) 20.
A species of the western half of the Amazonia, Swart-
zia bracteosa is now known, through the collection cited
below, well within Colombian territory. It is possible
that the type was from Colombia, for it was collected
by Martius ‘‘in sylva Yapurensibus. ”’
Cotompia: Intendencia del Amazonas, Rio Igaraparand, los alrede-
dores de La Chorrera, alt. ca. 180 m. ‘Small tree. Fruit red.”’
June 4-10, 1942, Richard Evans Schultes 3939.
VOCHYSIACEAE
Vochysia lomatophylla Standley in Field Mus.
Publ. Bot. 22 (1940) 150.
Vochysia lomatophylla was described from material
collected on the Rio Mazin in Loreto, Peru; it has also
been known from the Rio Solimdées in Brazil (Stafleu in
Rec. Trav. Bot. Néerl. 41, bk. 2 (1948) 505). The col-
lections cited below are the first recorded from Colombia
and greatly extend the known distribution of the species
to the headwaters of the Rio Negro basin.
Cotompta: Comisaria del Vaupés, Upper Vaupés River, alt. about
300 m. (?) January, 1944, Gabriel Gutiérrez & Richard Evans Schultes
504.—Comisaria del Vaupés, Upper Vaupés River, near Miraflores,
alt. about 300 m. (?) ‘‘Enormous tree on floodland along banks.’’
January, 1944, Gabriel Gutiérrez & Richard Evans Schultes 701.
Vochysia obscura Warming in Martius FI. Bras.
18, pt. 2 (1875) 73, t. 13.
Widely distributed in the Amazon basin of Brazil and
Peru and in southern Venezuela, this species of savannas
and caatinga-forests has not hitherto been reported from
Colombia. The collection cited below extends the known
range very far to the northwest.
Cotompia: Comisaria del Vaupés, Cerro Chiribiquete, January,
1944, Gabriel Gutiérrez & Richard Evans Schultes 688,
[ 28 ]
Vochysia vismiifolia Spruce ex Warming in Mar-
tius Fl. Bras. 18, pt. 2 (1875) 99.
This species of savannas or open caatinga-like forests
is widely distributed throughout the Amazon Valley of
Brazil and in southern Venezuela, but has hitherto ap-
parently not been recorded from Colombia.
Cotomsra: Comisaria del Vaupés, Cerro Chiribiquete, January,
1944, Gabriel Gutiérrez & Richard Evans Schultes 638,
EUPHORBIACEAE
Euphorbia penicillata (Millspaugh) R.E. Schultes
comb. nov.
Tithymalus pencillatus Millspaugh in Field Mus.
Publ. Bot. 4 (1925) 95.
Although the binomial Huphorbia penicillata has been
used in several enumerations (Herrera: ‘‘Catdlogo alfa-
bético de los nombres vulgares y cientificos de plantas
que existen en el Perti’’ (1989) 192; Herrera: ‘‘Sinop-
sis de la flora del Cuzco’’ 1 (1941) 291), it seems that the
combination has never properly been made. The specific
epithet was originally published as pencillatus, an obvious
typographical error which was corrected in the index of
the volume.
Phyllanthus fluitans Bentham ex Mueller-Argo-
viensis in Linnaea 82 (1868) 36.
Described from Brazil, Phyllanthus fluitans has ap-
parently not hitherto been reported from the flora of
Colombia.
CotomsiA: Intendencia del Amazonas, near mouth of Rio Loreto-
yacu, April, 1944, Frederick J. Hermann 11298.
Peru: Departamento de Loreto, Rio Amazonas, Isla Cacao, April,
1944, Frederick J. Hermann 113144.
[ 29 |
SAPINDACEAE
Serjania dasyclados Radlkofer var. sibundoya
R. EE. Schultes var. nov.
Frutex scandens, a Serjania dasyclados principaliter
foliolis subtus densius tomentoso-pilosis, supra dense et
sordide adpresso-pilosis et tactu mollibus differt.
This concept would seem to represent either a south-
ern variant of Sexjania dasyclados or an endemic variety
possibly confined to the mountain-encircled Valley of
Sibundoy where a large number of endemic plants have
been found. It is at once distinguished from Serjania
dasyclados by having dense and appressed hairs of a dirty
brown color on the upper surface of the leaflets. The
more northern collections of the Colombian Serjania
dasyclados (the type and a cotype from Cundinamarca
at 700 and 1000 m. respectively; a collection from Cal-
das at 1800-2100 m.; one from Santander at 2000-2600
m.; one from Norte de Santander at 2000-2500 m.)
have no indumentum on the upper surface of the leaflets.
CotomsiA: Comisaria del Putumayo, Valley of Sibundoy, Sibundoy,
alt. about 2225-2300 m. ‘*Vine. Flowers white; fruits ripen red.
Hairs on both leaf surfaces.’’? May 29, 1946, Richard Evans Schultes
& Mardoqueo Villarreal 7652 (Tyre in Herb. Gray).
BoMBACACEAE
Quararibea putumayensis Cuatrecasas in Lloydia
11 (1948) 186.
This collection is topotypical. The species is known
only from the Comisaria del Putumayo and adjacent
regions of Peru. The Kofan Indians employ the roots
of this bush in preparing arrow-poison.
CotomsiA: Comisaria del Putumayo, Rio Putumayo, Puerto Ospina.
**Flowers white. Fruit edible. Bush. Kofan: hun-kun-ni-ai.’’ March
23-25, 1942, Richard Evans Schultes 3420,
[ 30 ]
Quararibea Schultesii Cuatrecasas sp. nov.
Frutex. Ramuli badii, glabri, sublaeves, novelli pal-
lide virides, pilis stellato-pubescentibus vestiti. Folia
grandia, membranacea, simplicia, alterna, petiolata, pal-
lide viridia. Petiolus robustus, 1.5—2 em. longus, teres,
dense stellato-tomentosus. Lamina obovata, plusminusve
oblonga, basim versus attenuata, basi cuneata, apice
subito contracta et cuspidata margine integro; tri-quin-
tuplicinervis, nervis secundariis quattuor, utroque latere
duo tertio superiores dispositis aseendentibus prope mar-
gine arcuato-anastomosatis reliquis nervis transversis et
reticulum formantibus; supra prospectu glabra sed pilis
sparsissimis simplicibus munita, nervis principalibus con-
spiculs, reticulo obsoleto; subtus pilis sparsis simplicibus
et stellatis praecipue ad nervos praedita, nervis primariis
secundariisque valde eminentibus, reticulo bene emi-
nenti; 18-30 cm. longa, 10-15 cm. lata. Stipulae lan-
ceolatae, 9 mm. longae, 2.5 mm. latae, dense stellato-
tomentosae, mox deciduae. Flores solitarii, oppositifolii,
pedunculati. Pedunculi mediocres, flexuosi, circa 18 mm.
longi, striolati, dense adpresse stellato-tomentosi, apice
tres bracteas, late lanceolatas, dense stellato-tomentulo-
sas, circa 1 cm. longas, 2—4.5 mm. latas, basi calycis ad-
natas ferentes. Calyx tubuloso-conicus; tubus 13 mm.
longus, basim versus minute striatus, apice breviter 5-
lobulatus, dorso alis quinque membranaceis, apicem ver-
sus dilatatis, extremis rotundatis, calycis tubum superan-
tibus instructus, extus tubo alisque stellato-tomentulosis
intus adpresse albido-sericeis. Petala 8 em. longa, supra
glabra, subtus villoso-lanata cum pilis stellatis radiis ten-
uibus longis intricatis, verosimiliter albida; ungui late
lineari, glabro, 16 mm. longo; limbo apice exserto,
spathulato, 14 mm. longo, 5-6 mm. lato, apice rotun-
dato, reflexo vel patenti. Staminorum columna quam
calyx plus duplo longior, glabra, cirea 3 cm. longa, ex-
[ 31 ]
trema in lacinias quinque antheriferas lineares, 4—5 mm.
longas producta; laciniis sex antherarum loculis, oblon-
gis, 1-1.2 mm. longis, valde contiguis munitis, et loculis
alteris (1-2) parte superiore tubo concrescentibus. Stylus
longe villosus, antherarum lobulis parum brevior vel ae-
quilongus. Stigma capitatum, 1.8 mm. crassum.
In connection with the foregoing description, Cuatre-
casas states: ‘‘Quararibea Schultesiu belongs to the small
group of species which have wings on the calyx. The
most similar to it is Quararibea bracteolosa (Ducke)
Cuatr. but Q. Schultesiu differs in having longer petioles
and pedicels, by its smaller flowers, by the shorter mem-
branaceous calyx of which the wings are more expanded
and even more membranaceous, by the herbaceous-
membranaceous lanceolate bracts, and by the indumen-
tum of calyx and bracts which is shorter and thinner.
Quararibea bracteolosa, on the contrary, has bracts which
are ovate and coriaceous or subcoriaceous, a calyx tube
which is much thicker and dense. I have seen an isotype
of Quararibea bracteolosa which is preserved at the
Chicago Museum of Natural History. Probably Ducke
1781 and 1782 (fruiting specimens), distributed as Quar-
aribea bracteolosa, do not belong to that species. ”’
Cotomsi1A: Comisaria del Amazonas, Trapecio amazénico, Loreto-
yacu River, alt. about 100 m. ““Bush.’’ October 20-30, 1945, Richard
Evans Schultes 66694 (Tyrer in Chicago Museum).
OcHNACEAE
Rhytidanthera magnifica (Gleason) Dwyer in
Lloydia 9 (1946) 53, t. 2, a-g.
In his monographic treatment of Rhytidanthera and
related genera, Dwyer (loc. cit.) reported but one collec-
tion of R. magnifica, the type, Lawrance 599, from the
El Umb6o region, Departamento de Boyaca.
Cuatrecasas 13370 extends the known range of Rhyti-
[ 32 |
danthera magnifica into the Departamento Norte de San-
tander and very near to the Venezuelan boundary. A|-
though the type was described as a tree up to 65 feet
tall, the species may also be, as reported by Cuatrecasas,
a small tree (‘‘arbolito’’). Dwyer states that Rhytidan-
thera magnifica ‘‘is readily distinguished from the two
other species. .. . by its narrower leaflets which taper to-
ward the base....’’, but the leaflets of Cuatrecasas 13370,
while very narrow, are broadly rotund at the base.
CoLtomsia: Departamento Norte de Santander, regién del Sarare,
hoya del rio Marque entre Junin y Cordoba, 920-1240 m. alt. “‘Ar-
bolito. Pétalos blancos. Estambres amarillos.’’? November 22, 1941,
J. Cuatrecasas 13370,
Rhytidanthera mellifera R. HL. Schultes sp. nov.
Arbor parva, usque ad quindecim pedes alta, ramosis-
sima. ‘Truncus tres ad quattuor pollices in diametro, rufo-
brunneo cum cortice ruguloso et minute lenticellato.
Folia imparipinnata, aliquid pendula; petioli 16.5-24.5
em. longi; foliola duodecim vel plerumque tredecim
(cum petiolulis 1-2 mm. longis), lateralia alterna, 2.2-2.5
em. distantia, anguste lanceolata, apice acuta, basi ro-
tundata, 8.5-14 (plerumque 10.5) cm. longa, 2.2-3.2
(plerumque 2.5) cm. lata; foliolum terminale liberum,
cuneato-lanceolatum, apice obtusum, 10-12 cm. longum,
usque ad 4 cm. latum; alia marginem versus valde re-
trocurvata, margine grossiuscule serrata, dentibus apice
incurvis et plerumque 6 mm. distantibus, supra nitida
venis non elevatis, subtus pallidiora, venis prominenter el-
evatis, secundariis plusminusve viginti; stipulis caducis.
Inflorescentiae terminales, parvae, quam folia multo
breviores, 20 cm. longae, rhachide 8 mm. in diametro,
pauciflorae (floribus usque ad viginti); rhachidis rami
usque ad 8 cm. longi; pedicelli ad rhachidem articulati,
robustiores et lignosi, usque ad 10 mm. longi, apice sub-
clavati, basi 1 mm. et apice 2 mm. in diametro. Flores
[ 33 ]
solitaril, aromatico-fragrantes, mucilaginosi, alabastris 16
mm. longis, 7-8 mm. latis. Sepala quinque, imbricata,
concava; duo exteriora subcoriacea sed marginem versus
membranacea, rotundata, margine irregulariter lacerata,
apice profunde retusa, supra minute pulverulento-strio-
lata (non strigillosa), 7 mm. longa, 7 mm. lata, basi cum
plusminusve quindecim glandulis minutis digitaliformi-
bus, usque ad 1 mm. longis, 0.25 mm. in diametro; in-
teriora majora, membranacea vel apicem versus aliquid
papyracea, oblonga, apice profundissime fissa, margine
integra (vel sepalo extimo minute lacerato), 18 mm. longa,
7 mm. lata. Petala quinque, alba, membranacea, leviter
inaequalia, oblongo-spathulata, apice saepe obliqua (ala-
bastrum aliquid subcuculliforme), margine integra, 17
mm. longa, 7 mm. lata. Stamina sexaginta quattuor,
brunneo-flava, subaequalia, usque ad 12 mm. longa. An-
therae 10 mm. longae, 0.8 mm. latae, longitudinaliter
rugulosae, filamentis 2 mm. longis, basi 0.6 mm. latis.
Ovarium crassum, nigrum, inverse claviforme, apicem
versus in stylum indeterminatum, apparenter erectum,
luteum, paulatim angustans, 183 mm. longum, 2.5 mm.
in diametro, quinque cum stigmatibus sessilibus. Capsu-
lae immaturae solum visae, usque ad 2 cm. longae, 2.5—
3 cm. latae, praecipue apicem versus anguste subfalcatae.
Rhytidanthera, closely allied to Godoya, is the only
compound-leaved genus of the Ochnaceae. Hitherto,
there have been but three species known: Rhytidanthera
magnifica, R. splendida and R. suleata. The genus is en-
demic to Colombia.
Rhytidanthera mellifera represents the first collection
of the genus from the Amazon Valley, the other species
being Andean in their distribution. This is an excellent
example of the curious type of plant distribution and
relationship of the plants on the are which forms the so-
called Venezuelan-Guianan land-mass. Of this distribu-
[ B34 |
tion, Dr. Lyman B. Smith (in Contrib. Gray Herb. no.
161 (1946) 81) has said: ‘‘....movement on the are
was from the Andean region eastward. Finally most of
the species [i.e. of certain bromeliads and cyperaceous
plants he was discussing ] are narrow endemics, indicating
that the movement was relatively ancient. ’”’
Rhytidanthera mellifera is an endemic of the cretaceous,
quartzite hills of the upper Apaporis basin. We might
well expect Rhytidanthera to occur in collections from
other sandstone mountains in eastern Colombia and
southern Venezuela, but even now we may venture to
say that it is not acommon element of the flora of these
repositories of endemics. On the Cerro Castillo, Rhyti-
danthera mellifera is extremely rare. Special search for
more abundant flowering material failed to furnish addi-
tional individuals, so the species is at present known only
from one bush. This bush, growing on the edge of a rock-
ledge covered with a scrubby vegetation, was visible to
us far below as we were climbing the face of the moun-
tain. Like the other species of the genus, it makes an
unusual display with the large, white caesalpinaceous-like
flowers. Rhytidanthera was not seen on the nearby moun-
tains, Chiribiquete and Campana.
Rhytidanthera mellifera, so named because of the sweet
and mucilaginous exudation of the flower which attracts
swarms of wild bees to the plant, is immediately set apart
from the other known species by having 64 stamens, a
much larger number than has hitherto been found in the
genus.
The closest ally of Rhytidanthera mellifera would ap-
pear to be BR. magnifica (Gleason) Dwyer, from which
it differs in having twelve or thirteen alternate (instead
of nine or ten opposite) leaflets ; a very broadly cuneate-
lanceolate, obtuse (instead of a narrowly lanceolate-
elliptic, acute) terminal leaflet; much smaller lateral
[ 35 ]
leaflets; minutely lacerate (instead of entire) sepals;
smaller outer and larger inner sepals; and generally
smaller petals which, in the bud, are subcucullate at the
apex. There are also minor floral and vegetative differ-
ences which more abundant material may show to be of
importance.
Cotomsi1a: Comisaria del Caquetaé, Apaporis River, Cerro Castillo
(below confluence of Ajaju and Macaya), quartzite base, alt. of col-
lections 350-1000 feet above the forest floor, 1250-1900 feet above
sea-level. ““Flowers white, centres yellow-brown, aromatic fragrant.
Treelet 15 feet. On exposed rock. Much branched. Basal diameter 3-
4 inches. Leaflets very coriaceous, marginally inrolled.’’ July 27,
1943, Richard Evans Schultes 5657 (Tyrer in U.S. Nat. Herb.).
Sauvagesia erecta Linnaeus Sp. Pl. (1753) 203.
Sauvagesia erecta, the most widespread species of the
genus, is rather variable and presents several general geo-
graphical variants. It has been collected in Colombia in
the Departamentos de Norte de Santander, Huila, To-
lima and Cauca (Dwyer in Bull. Torr. Bot. Club 72
(1945) 580). The collection cited below is apparently the
first from the entire Amazon basin of Colombia. There
are numerous collections from the adjacent Amazonian
regions of Brazil and Peru.
Cotomsia : Comisaria del Amazonas, Rio Igaraparanaé, near La Chor-
rera, alt. about 180 meters. ‘‘Small shrub under 8 inches tall. Flow-
ers pinkish.’’ June 4-10, 1942, Richard Evans Schultes 3955.
FLACOURTIACEAE
Lindakeria nitida Willip & R. I. Schultes sp. nov.
Arbuscula usque ad viginti quinque pedes alta, ramo-
sissima, ramis longis laxisque. Folia ovata, basi cuneato-
obtusa, apice attenuato-acuminata, integerrima, 10.5—14
cm. longa, 4.5—5.5 em. lata, supra nitidissima, vivo ap-
parenter atroviridia cum venis non conspicuis, subtus
aliquid pallidiora cum venis prominentibus, utrinque
glaberrima, nervis secundariis plusminusve quinque. Pet-
[ 36 ]
ioli 2.5-8.5 em. longi, teretiusculi, ad laminam incrassati.
Racemi axillares, apparenter erecti et stricti; rhachides
usque ad 10 cm. longae, folia subaequantes, compresso-
angulatae. Pedicelli distichi, 6-7 mm. longi, solitarii vel
saepe duo vel tres. Flores 10 mm. in diametro vel mi-
nores, alabastro globoso. Sepala tria, triangularia, apice
subacuta, margine valde involuta, 4.5 mm. longa, basi
4 mm. lata, extus apparenter minute aspero-pustulata et
basi pilosa. Petala sex, alba, membranacea, lanceolata,
apice acuta, 7 mm. longa, 1.8 mm. lata, extus rarenter
dense albido-leprosa. Stamina plusminusve quadraginta,
4—5 mm. longa; filamenta libera, laxa, aliquid compla-
nato-filiformia, 2 mm. longa; antherae lineares, 2.5-8
mm. longae. Ovarium ovoideum, echinato-tuberculatum,
2 mm. in diametro. Stylus ovario circiter duplo longior,
3.5-3.8 mm. longus, complanato-filiformis, glaber, stig-
mate minute trilobo. Fructus adhue ignotus.
Lindakeria nitida is at once distinguished from all
other species of the genus by its very long and lax inflo-
rescence which gives the plant a completely unique as-
pect. This species, growing in the Magdalena watershed
of Colombia, is perhaps most closely allied to the rather
widespread Amazonian Lindakeria maynensis. In addi-
tion to the length of its inflorescence and its highly lus-
trous leaves, Lindakeria nitida differs from L.maynensis
in the shape and size of the sepals and slightly in the
structure of the stamens.
The collection Haught 1917 has, in general, much
larger leaves and flowers than the type. The leaves of
this cotype are lanceolate-ovate, up to 26 cm. long and
8 cm. wide; the petioles measure up to 7 em. in length,
and the rachis is 16 ecm. long. Additional collections will
be needed before we can determine whether the type
represents the normal condition of the species, but it
would appear from an examination of the available ma-
[ 37 |
terial that Haught 1917 was taken from an individual
in an open, favorable site or else from a new flush.
Cotompia: Departamento E] Magdalena, 17 km. north of El Banco
on road to Chimichagua. “‘Large (5 m. high) shrub with many long,
lax branches. Inflorescence very conspicuous. Corolla white, anthers
yellow. Inswampy forest, along road.’*? August 3, 1938, Oscar Haught
2221 (Tyre in U.S. Nat. Herb. No. 1706900).—Departamento de
Santander, El Centro, vicinity of Barranca-Bermeja, Magdalena Val-
ley, between Sogamoso and Carare Rivers, alt. 100-500 meters. ‘‘Large
shrub (4 m.) very conspicuous white flowers and buds.’’? August 26,
1936, Oscar Haught 1917.
Mayna glomerata Killip & R. EH. Schultes sp. nov.
Arbor parva, usque ad duodecim pedes alta. Rami
ramulique verrucoso-striati, griselli; ramuli juveniles
cinereo-virides, squamiferi. Folia alterna, simplicia, lan-
ceolato-elliptica, usque ad 22 cm. longa (sine petiolo),
7-7.5 cm. lata, integra, apice longe acuminata, basi cu-
neata, chartacea, nervis secundariis valde arcuatis duo-
decim vel tredecim, omnino glabra, supra atroviridia et
apparenter nitida cum nervis leviter prominentibus sed
non elevatis, infra pallide viridia, non nitida, nervis valde
elevatis et fulvis. Petioli robusti, usque ad 8 cm. longi,
1.5-1.8 mm. in diametro, glabri. Stipulae non visae.
Dioecia. Flores in fasciculis caulinis, glomeratis, inflores-
centiis usque ad plusminusve decemfloris. Alabastra glo-
bosa vel elongato-globosa, plusminusve 2 mm. in diam-
etro, minute et subdense pilosa. Bracteae minutae. Pe-
dunculus usque ad 8 mm. longus, dense albo-pilosus.
Flores staminiferi albi, conspicui, usque ad 2 em. in di-
ametro. Sepala tria, leviter inaequalia, valde concava,
ovata, integra, apice subacuta, extus pubescentia, intus
glabra, quinque cum venis brunneis longitudinaliter stri-
ata, sepalum majus 3.8-4 mm. longum, 2.7-2.9 mm.
latum. Petala quinque ad septem, aequalia, subspathu-
lata, margine irregularia, apice late rotundata et valde in-
voluta, alba, membranacea, usque ad 6.5 mm. longa, apice
[ 38 |
circiter 2 mm. lata, basi 0.4 mm. lata, omnino glabra.
Stamina circiter vigintiquinque, 2.6 mm. longa, filamen-
tis liberis, erectis, plerumque 0.8 mm. longis, et antheris
leviter arcuatis, 1.8 mm. longis, 0.5 mm. latis, flavis,
apice poricidis. Fructus adhuc ignotus.
Mayna glomerata is easily distinguished from the
other species of this genus by its unusual inflorescences
which are in small, closely packed, glomerate fascicles.
There are other notable differences, as well, such as the
curious folding of the upper third of the petals, the shape
of the petals, and the number of the stamens.
Cotompra: Departamento de Santander, vicinity of Barranca Ber-
meja, Magdalena Valley, between Sogamoso and Carare Rivers, Vis-
cania Creek, 32 km. south of El Centro, at about 100 meters alt.
**Small tree, about 4 m. high. Flowers white with yellow centres,
showy, closely covering twigs.’’ November 14, 1936, Oscar Haught
2070 (Typr in U.S. Nat. Herb. No. 1742387).
Mayna longifolia Poeppig & Endlicher Nov. Gen.
ac Sp. 3 (1845) 64, t. 271.
Mayna longifolia is extremely abundant in the forests
of the southern half of the trapecio in areas above flood-
level. The collection Llewelyn Williams 2711 from La
Victoria (now ‘‘Granja Caldas’’) near Leticia, cited as
from Peru in Macbride’s ‘‘ Flora of Peru’’ (Field Mus.
Publ. Bot. 18, pt. 4, (1941) 18), but actually from Co-
lombian territory, is referable to Mayna longifolia.
CoLoms1a: Comisaria del Amazonas, Trapecio amazénico, Loreto-
yacu River, alt. about 100 m. ““Treelet 25 feet.’’ October 1945,
Richard Evans Schultes 6700.—Comisaria del Amazonas, same locality
and date. ‘“Tall treelet. Leaf larger than normal.’’ Richard Evans
Schultes 6701.—Comisaria del Amazonas, same locality and date.
“Leaves very narrow. Fruit on trunk up to top.’’ Richard Evans
Schultes 6705.
COMBRETACEAE
Combretum laxum Jacquin Enum. Pl. Carib. 19
(1760) 115.
[ 39 |
Combretum laxum appears to be an aggregate species
in which several clearly distinct ‘‘tendencies’’ can be
seen. As now understood in this broad sense, the species
occurs all over tropical America. Several collections from
the Departamento de Loreto, Peru, represent the same
variant as Schultes 3924 from adjacent Colombia. When
further material is studied, this variant can probably be
shown to be specifically distinct from true Combretum
laxum.
In the Igaraparana, the Witoto Indian name of Com-
bretum lawum is qjuo-wo. In another Witoto dialect from
Loreto, Peru, the native name has been recorded as
aioho-ayo-o (in Field Mus. Publ. Bot. 18, pt. 4 (1941) 225).
This attractive vine often covers riverside vegetation
and, especially at night, gives off an extraordinarily
strong and sweet perfume.
Cotompra: Comisaria del Amazonas, Rio Igaraparana, alrededores
de La Chorrera. “‘Vine. Flowers yellow, fragrant.’’ June 4-10,
1942, Richard Evans Schultes 3924.
MELASTOMACEAE
Aciotis aequatorialis Cogniaux in Martius FI.
Bras. 14, pt. 8 (1885) 464.
Known from the Pacific coast of Colombia, from An-
tioquia and from the Intendencia del Meta, Aciotis ae-
quatorialis has hitherto apparently not been reported as
an element of the flora of Amazonian Colombia. A ciotes
polystachys (Bonpl.) Tr. and A. purpurascens (Aubl.)
Tr. have been collected in Colombia only a few kilo-
meters from the locality of the specimen of 4. aequa-
torialis cited below.
CoLomspia: Intendencia del Amazonas, Rio Hamacayacu, April,
1944, Frederick J. Hermann 11303.
Leandra divaricata (Naud.) Cogniaux in Martius
Fl. Bras. 14, pt. 4 (1886) 196.
[ 40 |
Not infrequent in British Guiana, Venezuela, Ama-
zonian Peru and elsewhere in Colombia, Leandra divari-
cata has been collected hitherto on the Amazonian water-
shed of Colombia but twice: at Puerto Porvenir, on the
upper reaches of the Rio Putumayo (Cuatrecasas 10666),
and at Mocoa (Cuatrecasas 11316). Hermann 11308,
from the southeasternmost point of the Colombian Am-
azonia, would seem to indicate a wide distribution for
this species in eastern Colombia.
CoLompia: Intendencia del Amazonas, Rio Hamacayacu, April,
1944, Frederick J. Hermann 11308.
LOGANIACEAE
Strychnos amazonica Krukoff in Brittonia 4
(1942) 284.
This collection, determined with some reservation as
representing Strychnos amazonica, is the first of the spe-
cies from Colombia. Krukoff and Monachino (in Caldasia
4 (1946) 45) enumerated Strychnos amazonica as a species
to be expected in the Colombian Amazonia. It is one of
the plants the root of which is used by the Kofin Indians
in preparing arrow-poison.
CoLtomsia: Comisaria del Putumayo, Rio Putumayo, Puerto Ospina.
6 . . r r ‘ . ,
‘Vine. Root used in Kofan Indian arrow-poison. Kofan name: hess-
pd-chu.”’’ April 25, 1942, Richard Evans Schultes 3689.
Strychnos Erichsonii Richard Schomburgk Faun.
Fl. Br. Guiana (1848) 1082, nomen: ex Progel in Mar-
tius Fl. Bras. 6, pt. 1 (1868) 274.
First reported from Colombia on the basis of a collec-
tion from the Rio Sucumbios (in Bot. Mus. Leafl. Har-
vard Univ. 13 (1949) 290), Strychnos Erichsonii is appar-
ently one of the most abundant species of this genus in
the upper Putumayo basin where it is sought by the
Kofan Indians for use in preparing cuvrare.
[ 41 ]
Cotomsia: Comisaria del Putumayo, Rio Putumayo, Puerto Ospina.
“*Vine. Bark of stem rasped and used for arrow-poison. Kofin name:
ee-ru-che.’’ April 25, 1942, Richard Evans Schultes 3682.—Same lo-
cality and date. ‘*Vine. Root used in arrow-poison, Kofaén name:
see-a-se-sep.’’ Richard Evans Schultes 3682.—Same locality and date.
““Vine. Pieces of stem used in preparing Kofan arrow-poison. Kofan
name: mun-tai-ri-chee.’’ Richard Evans Schultes 3685.
Strychnos guianensis (4ub/.) Martius Syst. Mat.
Med. Bras. (1848) 121.
Known from Colombia previously from one collection
made in the Rio Vaupés, Strychnos guianensis is now
registered from the southwesternmost point of the Am-
azon basin of Colombia where it serves the Kofin Indians
as one of the lesser ingredients in arrow-poisons.
Cotomsi1A: Comisaria del Putumayo, Rio Putumayo, Puerto Ospina.
ee e . . 4 , 99
Vine. Root used in preparing curare. Kofin name: ku-see-ye-hé-pa.
April 25, 1942, Richard Evans Schultes 3688.
Strychnos javariensis Avrukoff in Brittonia 4
(1942) 279.
The following collections are apparently the first to be
recorded from Colombia. The species is one of the lesser
ingredients of Kofan curare.
Cotompra: Comisaria del Putumayo, Rio Putumayo, Puerto Ospina.
‘*Vine. Root used in arrow-poison. Kofan name: tzu-se-hé-pa.’’ April
25, 1942, Richard Evans Schultes 8690.—Same locality and date.
**Vine. Root used in Kofan arrow-poison.’’ Richard Evans Schultes
3691.
Strychnos Jobertiana Bail/on in Adansonia 12
(1879) 367.
Although previously reported from two localities in
the Colombian Amazonia, Strychnos Jobertiana is now
known to be very abundant in the upper Putumayo
basin and to be one of the preferred ingredients of curare
amongst the Kofan Indians.
(42)
Cotomspia: Comisaria del Putumayo, Rio Putumayo, Puerto Ospina.
**Vine. Stem employed as ingredient in arrow-poison. Kofaén name:
u-su-se-é-pa.’” April 25, 1942, Richard Evans Schultes 3692.—Same
locality and date. ““Vine. Root used to make arrow-poison. Kofan
name: fee-see-pd-chu.’’ Richard Evans Schultes 3695.
Strychnos Peckii B. L. Robinson in Proc. Am.
Acad. 49 (1918) 504.
This is the second collection of Strychnos Peckii from
Colombian Amazonia. It is one of the preferred species
amongst the Kofan Indians for preparing arrow-poison.
Cotoms1a: Comisaria de] Putumayo, Rio Putumayo, Puerto Ospina.
“Vine. Root used in arrow-poison. Kofan name: su-he-sé-pa.’’ April
25, 1942, Richard Evans Schultes 3684.
Strychnos subcordata Spruce ex Bentham in
Journ. Linn. Soc. 1 (1856) 106.
This species has never been reported from Colombia.
It is considered one of the strongest of the curare ingre-
dients by the Kofan Indians.
Cotoms1A: Comisaria del Putumayo, Rio Putumayo, Puerto Ospina.
“*Vine. Root used in arrow-poison formula. Kofaén name: su-sé-pe.”’
April 25, 1942, Richard Evans Schultes 3687.
APOCYNACEAE
Himatanthus articulata (Vah/) Woodson in Ann.
Missouri Bot. Gard. 25 (19837) 196.
The collection cited below is apparently the second
from Colombia, the first being Cuatrecasas 9001 from
Florencia, capital of the Comisaria del Caquetaé. The
species is rather widespread in the Guianas, Amazonian
Brazil, the Orinoco drainage-area of Venezuela and in
Middle America.
. , , oe oe | 6é
CotompBia: Comisaria del Vaupés, Cerro Chiribiquete. Flowers
white.’’? January, 1944, Richard Evans Schultes 5735,
[43]
Himatanthus bracteata (4.DC.) Woodson in
Ann. Missouri Bot. Gard. 25 (1937) 200.
Known from British Guiana and several stations in
the Amazon of Brazil (where it is apparently most abun-
dant in the Rio Negro basin) as well as from Pernam-
buco and Bahia, Himatanthus bracteata has hitherto not
been reported from Colombia. Schultes 5814, tentatively
determined as representing this species, has, according to
Woodson, leaves which ‘‘are larger than usual.’’ In the
Vaupés, the plant is known as platanote, due to the
superficial resemblance of the fruit to the banana.
Cotomsia: Comisaria del Vaupés, Cerro Chiribiquete. ‘‘Flowers
white.’’ January, 1944, Richard Evans Schultes 5738.—Comisaria del
Vaupés, Rio Vaupés, Cafio Pact (below Cerro Circasia). ‘‘Small tree
up to 15 feet tall. Latex abundant, white. Platanote.’’ March 6,
1944, Richard Evans Schultes 5814.
Mandevilla nerioides Woodson in Ann. Missouri
Bot. Gard. 28 (1941) 274.
Mandevilla nerioides is known only from the Inten-
dencia del Meta and the Comisaria del Vaupés in Colom-
bia. The type collection was made on the Guaviare River
at San José del Guaviare by Cuatrecasas. The following
collections establish the occurrence of this endemic spe-
cies on the summit of the remnant quartzite hills of the
upper Apaporis River which have, in common with the
hills at San José del Guaviare, a number of. striking
endemic elements.
CotomB1a: Comisaria del Vaupés, Cerro Chiribiquete. ““Vine. Flow-
ers pink. Latex white, sticky.’’ May 15-16, 1943, Richard Evans
Schultes 5442.—Same locality. “*Vine. Flowers pinkish white. Latex
white.’’ January, 1944, Richard Evans Schultes 5740.
SOLANACEAE
Solanum quitoense Lamarck Llustr. 2 (1798) 16.
Although the /u/o, the fruit of this species, is known
[ 44 ]
in many parts of southern Colombia and has been the
subject of several recent articles (Chalons in Agric.
Amer. 4 (1944) 110-112; McCann, ibid. 7 (1947) 146-
149; Hodge in Rev. Fac. Nac. Agron. 7 (1947) 147-
154; Hodge in Journ. N.Y. Bot. Gard. 48 (1947) 155-
159), there are apparently no Colombian collections of
Solanum quitoense in the larger herbaria of Colombia and
the United States.
The lulo is used as a fruit and is cultivated in much of
Andean Colombia (Hodge, loc. cit.), especially in the
southern part of the country—in El Valle, Cauca and
Narifo (Pérez-Arbelaez, ‘‘Plantas medicinales y venen-
osas de Colombia’’ (1987) 246; ‘‘Plantas ttiles de Colom-
bia’? (1947) 451), where it is known also as naranjillo and
toronja. Lacking specimens or a critical taxonomic study
of the lulo from the numerous areas of Colombia, we
cannot be certain that only one species is involved.
Solanum quitoense, as represented in our herbaria, is
rather common in the Andean highlands of Ecuador and
Peru. The collections Schultes & Villarreal 7616 and
Schultes 3271 from the southernmost Andes of Colombia
not far from the boundary of Ecuador are taxonomi-
cally indistinguishable from the material from Ecuador
and Peru. It is obvious that an extensive taxonomic
study of the lulo in Colombia is needed. Certainly the
possibilities of this delicious fruit would indicate the ad-
visability of such an investigation.
Co.tomsra: Comisaria del Putumayo, Valley of Sibundoy, Sibundoy,
alt. about 2225-2300 m. “‘Large shrub. Stem and under surface of
leaf purplish, hairy. Flowers white, anthers yellow. Fruit orange
colored and size of oranges. Naranjillo. Kamsa Indian name: ma-sha-
kve.’’ May 29, 1946, Richard Evans Schultes & Mardoqueo Villarreal
7616.—Same locality, February 16, 1942, Richard Evans Schultes
3271.—Departamento de Antioquia, Rio Negro, ‘“La Granja.’’ Cul-
tivated, November 2, 1946, W. H. Hodge 67172.
[ 45 ]
RUBIACEAE
Rondeletia rupicola R. Schumann var. chiribi-
quetana Rk. EH. Schultes var. nov.
Frutex a Rondeletia rupicola stipulis foliaceis multo
minoribus; foliis elliptico-lanceolatis (non longe lanceo-
latis), minoribus, 4-5 cm. longis, 1-1.3 em. latis (non
5-8 cm. longis, 1.5 cm. latis), acutis (non acutissimis,
mucronatis), supra prominentius bullatis, prope nervos
majores densius sericeo-strigillosis, subtus prope nervos
atque in lamina ipsa maxime densissime sericeo-strigillo-
sis; corollae laciniis majoribus, extus densius strigillosis ;
antheris majoribus; fornicibus papillosis majoribus den-
sissime hirsutis differt. Capsula globosa, coriacea, 2mm.
in diametro, in fructu calycis laciniis usque ad 1.5 mm.
longis densissime strigillosis coronata, fusco-brunnea;
semina numerosissima, minuta, angulata,
A study of the original description of Rondeletia rupi-
cola and an examination of a duplicate type (Spruce 3392)
preserved at the Gray Herbarium and a photograph of
the Spruce specimen at Berlin indicate that there are a
number of minor variations which serve to separate the
new variety described above. The capsule of Rondeletia
rupicola is unknown, but the type of var. chiribiquetana
has both flowers and fruit. The color of the flowers of
Rondeletia rupicola is not designated; that of var. chiri-
biquetana is pink.
Cotomsia: Comisaria del Vaupés, Macaya-Ajaju River confluence,
Mount Chiribiquete, quartzite base, summit 800-1200 feet above
forest floor, 1700-2100 feet above sea level. ‘‘Flowers rose-pink.
Leaves dark green above, pale below. Hair silvery. Shrub.’’ May
15-16, 1943, Richard Evans Schultes 5448 (Tyrr in Herb. Gray).
COMPOSITAE
Clibadium asperum (4 1b/.) de Candolle Prodr. 5
(1836) 506.
[ 46 ]
This species of Clibadiwm is abundantly cultivated in
the vicinity of La Pedrera and in the Rio Miritiparana
for use as a fish-poison. It is known locally as guaco by
the Spanish-speaking inhabitants and as /v-na-pe by the
Mirafia Indians. Several other species of Clibadium are
widely cultivated in the Amazon regions as fish-poisons.
CoLtompia: Comisaria del Caqueté, Rio Caquetdé, vicinity of La
Pedrera, April, 1944, Richard Evans Schultes 5879.
Gongylolepis maroana Badillo in Bot. Soc. Venez.
Ciéne. Nat. 8 (19438) 287.
Recently, two stations for this species were reported
for Colombia, both in the Comisaria del Vaupés (in Bot.
Mus. Leafl. Harvard Univ. 13 (1949) 810). The collec-
tion cited below is the third from Colombia and extends
the known range of the plant to the west on the same
interrupted sandstone hills on which previously reported
collections were made.
Cotomsp1a: Comisaria del Vaupés, Ajaju River, Cerro de La Cam-
pana, quartzite base, summit about 800-1200 feet above forest floor,
1700-2100 feet above sea-level. ‘‘Composite shrub. Common on top
of mountain,’’ June 1-6, 1943, Richard Evans Schultes 5568.
[ 47 ]
ILLUSTRATIONS
EXPLANATION OF THE ILLUSTRATION
Pirate VII. (Upper figure). Habit photograph of
CrcROPIA PORVENIRENSIS Cuatr., Puerto Porvenir,
Rio Putumayo, Colombia.
(Lower figure). Flowering branch of Vocuysta Lo-
MATOPHYLLA Stand/, the tree from which Gutiérrez
& Schultes 701 was collected, Rio Vaupés, Colombia.
Photographs by R. E. Scuutters
Pruare VII
EXPLANATION OF THE ILLUSTRATION
Puare VILE. Ruyrrpanrurra Meviirera PR. FE,
NSchultes. 1, leaf one half natural size. 2, inflores-
cence, one half natural size. 3, flower, natural size.
4, stamen, about four times natural size. 5, anther
tip showing apical pores, about twenty times nat-
ural size.
Drawn by KE. W. Suivi
PLaTE VIII
mellifera
RFE. Schuttes
EXPLANATION OF THE ILLUSTRATION
Pirate IX. Linpakerta nitipa Killip & Schultes. 1,
flowering branchlet (Haught 2227), one half natu-
ral size. 2, budding branchlet (Haught 1719), one
half natural size. 3, flower, about two and one half
times natural size. 4, stamen, about five times nat-
ural size.
Drawn by EK. W. Suiru
PuaTE IX
LINDAKERIA
nitida
Fal lip &
Schultes
Q of,
)
Qu.
Qy TY
7 A
EXPLANATION OF THE ILLUSTRATION
Piare X. (Upper figure). Soranum qurromnse Lam.
Photograph of the plant from which Schultes & Vil-
larreal 7616 was collected.
(Lower figure). An escarpment of Cerro del Cas-
tillo, Rio Apaporis, Colombia. RuyTIpANTHERA
MELLIFERA Occurs on these open, exposed quarzitic
escarpments.
Photographs by R. F.. Scuu.tres
PLATE X
EXPLANATION OF THE [TLLUSTRATION
Prare XI. Mayna aromerata Killip & Schultes. 1,
Howering branchlet, one half natural size. 2, flow-
er, about three times natural size.
Drawn by Kb. W. Suivi
PLaTE XI
y MAYNA glomerata
v Lillip & Schultes
;
7’, ee ee ee ae, eee A + ee eee eS ee ee eee pee eee
BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY
VoL. 14, No. 3
CamsripGe, Massacuusetts, DecemMBer 7, 1949
NOMENCLATURAL NOTES
AND NEW CONCEPTS OF TROPICAL
AMERICAN ORCHIDS
BY
CHARLES SCHWEINFURTH
Durinec the course of recent studies, I have found it
necessary to make numerous changes in the status of
published concepts, to amplify certain descriptions and
to describe several orchids which appear to be new. This
addition to the number of recorded species and varieties,
together with many similar ones, shows that tropical
America from the northern boundary of Mexico to the
southern borders of Peru and Brazil is a veritable store-
house of plants new to science. Only the collector and
the taxonomist are necessary to bring these to light.
The arrangement of the species follows the order pro-
posed by Dr. Rudolf Schlechter in ‘‘Notizblatt des Bo-
tanischen Gartens u. Museums Berlin-Dahlem 9 (1926)
563-591.
Cranichis ciliilabia C. Schweinfurth sp. nov.
Herba terrestris, gracilis. Folia plura, plerumque ba-
salia et petiolata; lamina ovata, acuta vel breviter acu-
minata, parva. Caulis basi glaber, supra dense glanduloso-
pubescens, vaginis pluribus remotis ornatus. Inflorescen-
tia laxe pluriflora cum floribus parvis membranaceis.
Sepala elliptica vel ovalia, acuta. Petala oblique elliptica,
pro genere lata, dense ciliata. Labellum simplex, cuneato-
[ 49 ]
obovatum, antice late rotundatum, concavum, dense
longe ciliatum. Columna perbrevis.
Plant terrestrial, slender, about 21.5 cm. tall. Roots
tuberous, fasciculate, slender, lanuginose. Leaves several,
the four basal blades rosulate, slender-petioled; lamina
ovate, acute or short-acuminate, broadly cuneate to sub-
truncate at the base, commonly 3.2-3.5 cm. long, up to
2.2 cm. wide; petiole channelled, slightly dilated above
and below, about 2 cm. or less long. A fourth similar
but sessile leaf rises near the base of the stem. Stem
slender, glabrous in the lower portion, densely glandular-
pubescent above, bearing four distant sheaths of which
the lower ones have a close long-sheathing tubular base
and the lowermost is dilated into a small foliaceous blade.
Raceme loosely 17-flowered, about 4.5 cm. long, with
the apical portion (bearing immature flowers) nodding.
Flowers small, but medium-sized for the genus, mem-
branaceous. Sepals very sparingly glandular-pubescent
on the outer surface. Dorsal sepal elliptic, acute, con-
cave, 3-nerved with the lateral nerves branching below,
about 5.9 mm. long and 8.2 mm. wide when expanded.
Lateral sepals oval, acute, concave, 5-nerved, about 5.6
mm. long and 8.2 mm. wide. Petals obliquely elliptic,
acute or subacute, 4-nerved, densely ciliate with the cilia
longer on the anterior margin, about 5 mm. long and
2.7 mm. wide. Lip simple, cuneate-obovate, broadly
rounded in front, minutely acute at the apex, densely
long-ciliate, about 4 mm. long and 3 mm. wide near the
front in natural position, rather deeply concave especially
above, 3-nerved, the lateral shorter nerves recurved in
the middle of the lamina and having two or three short
recurved branches. Column very short, with the stalk
of the anther arising near the base.
The long-ciliate lip seems to be unique in the genus
and the ciliate petals are remarkably broad.
[ 50 |
Mexico: Chiapas, at 4000-6000 feet altitude, “‘moist bank of trail
above Liquidambar,’’ flowers white, November 9, 1945, A. J. Sharp
45977 (Tyrer in Herb. Ames No. 65446).
Pleurothallis Broadwayi 4 mes Orch. 2 (1908) 267.
Pleurothallis guadalupensis Cogniaux in Urban Symb.
Antill. 6 (1909) 432.
Pleurothallis Williamsti Ames Orch. 7 (1922) 120.
Pleurothallisnana A. & 8. in Sched. Orch. 8 (1925) 29.
The above synonymy, except for Pleurothallis nana,
has already been published (C.Schweinfurth in Bot. Mus.
Leafl. Harvard Univ. 8 (1940) 41).
At the time when Pleurothallis nana was erected, it
was thought to be distinct from the concepts referable
to P. Broadwayt. It seemed to have longer and more
branched stems than the other species of this group. It
also appeared to be separable by reason of its longer pedi-
cels, ‘‘snow white’’ rather than yellow or greenish yellow
flowers and especially because of its narrower and more
acuminate sepals.
A recently acquired Mexican collection (Nagel &
Juan G. 6447), although it appears to be inseparable
vegetatively from the Costa Rican P. nana, has long
pedicels and sepals intermediate in form between those
of typical P. Broadwayi and P. nana. It is further noted
as having pale yellow-green flowers, as in P. Broadway.
A Costa Rican collection (Austin Smith H541) also has
pedicels of a length intermediate between the two con-
cepts and flowers noted as pale lemon yellow as in P.
Broadway; but it has sepals of quite the form of P. nana.
Therefore, in view of the great variability naturally to
be expected of a wide ranging plant, it seems advisable
to regard P. Broadwayi as representing one polymorphic
species. It occurs in Mexico (Oaxaca), Honduras, Costa
Rica (type of P. nana), Panama (type of P. Williamsii),
[ 51 ]
Cuba, Martinique, Guadeloupe (type of P. gwadalupen-
sis) and Venezuela (Island of Margarita).
Pleurothallis leucantha Schlechter in Fedde Re-
pert. 10 (1912) 353.
Pleurothallis Sanchot Ames in Sched. Orch. 4 (May
1923) 26.
Pleurothallis gonioglossa Schlechter in Fedde Repert.
Beih. 19 (November 1928) 189.
Judging from the type description of the Guatemalan
Pleurothallis leucantha (amplified by an analytical draw-
ing made under the supervision of Dr. Schlechter), that
species differs from the Costa Rican P. Sanchoi (repre-
sented by the type and several subsequent collections),
by having generally longer and more acuminate leaves,
smaller flowers, somewhat dissimilar straighter petals and
an obtuse (not apiculate) lip.
In the Ames Herbarium there is a Guatemalan collec-
tion (Harry Johnson 609) taken to represent P. leucan-
tha which came from the same region of Coban as the
type and at about the same altitude. This collection,
which may be called atopotype of P./eucantha, has some-
what larger flowers than specified as typical, slightly
curved petals and an apiculate lip which very nearly
coincides with that of P. Sanchot. Moreover, one Costa
Rican collection (Standley & Valerio 45983) is entirely
typical of P. Sanchoi, except that the petals are nearly
straight, as in typical P. leucantha.
The concept described as Pleurothallis gonioglossa,
which is represented by an isotype and analytical draw-
ings in the Ames Herbarium, appears to be an exact
vegetative counterpart of P. Sanchoi, although it has
slightly larger flowers than the latter species. Except
for inconsequential floral differences, this species appears
to be inseparable from P. leucantha.
[ 52]
It seems logical, therefore, to regard Pleurothallis leu-
cantha as a rather widespread and variable species in
Central America.
Octomeria pygmaea C. Schweinfurth sp. nov.
Herba minutissima, epiphytica, corticicola. Caules
perbreves, uniarticulati, vaginis duabus membranaceis
tubulatis evanidis omnino obtecti. Folium anguste ellip-
ticum, acutum, sessile, rigide nervosum, crasse margina-
tum. Inflorescentiae saepissime duae vel tres,abbreviatae,
uniflorae. Flos membranaceus, perparvus sed pro planta
magnus. Sepala persimilia, triangulari-lanceolata, acuta
vel acuminata, trinervia, omnino libera. Petala lanceo-
lata vel elliptico-lanceolata, acuta vel acuminata, uni-
nervia. Labellum in circuitu rhombico-ovatum, medio
profunde trilobatum ; lobi laterales faleato-dolabriformes,
porrecti; lobus medius obovato-oblongus, late obtusus,
lobos laterales valde excedens. Columna in pedem sub-
aequalem extensa.
Plant minute, caespitose, epiphytic in the cracks of
the bark. Roots fibrous, relatively stout, glabrous. Stems
numerous, clustered, abbreviated, about 4.8 mm. or less
long, 1-jointed near the base, entirely concealed by two
tubular scarious evanescent sheaths. Leaf solitary, ses-
sile, narrowly elliptic, up to about 8 mm. long and 2.1
mm. wide, acute, conduplicate at the base, coarsely mar-
ginate, rigid-nervose, with the mid-nerve enlarged and
prominent beneath. Inflorescences abbreviated, 1-flow-
ered, apparently two or three to a stem. Flower very
small, but large in relation to the plant, membranaceous.
Sepals free, 3-nerved, concave. Dorsal sepal triangular-
lanceolate, acute or acuminate, about 1.75 mm. long and
0.75 mm. wide. Lateral sepals very similar, but a little
longer and wider, oblique. Petals lanceolate to elliptic-
lanceolate, acute or acuminate, 1-nerved, about 1.5 mm.
[ 58 ]
long and 0.5 mm. wide. Lip rhombic-ovate in outline,
deeply 8-lobed near the middle, cuneate at the base,
about 1.1 mm. long and 0.87 mm. wide across the lateral
lobes, with a pair of fleshy keels through the lower por-
tion; lateral lobes faleate-dolabriform, subequal to the
mid-lobe, obtuse, porrect, erect in natural position; mid-
lobe obovate-oblong, broadly obtuse, much exceeding the
lateral lobes. Column about half as long as the petals,
produced into a subequally long foot. Pollinia eight,
pyriform.
This species is probably the most minute member of
the genus Octomeria yet described and ranks with the
most insignificant orchids known to science. It appears
to be allied to O. minuta Cogn., but differs in having
acute sepals (when expanded), the lateral ones being free,
as well as in having a widely dissimilar lip.
I take the liberty of quoting from the notes of the
collector of this remarkable little orchid, as follows:
‘*A fter making camp one afternoon I put on the climb-
ing irons in order to collect a specimen of a beautifully
flowering Cunuria crassipes, a relative of Hevea rubber.
When half way up the ninety foot tree, I stopped to
lean back in the belt and rest. My glance was caught by
what seemed to be a tiny orchid growing, together with
the ever-present mosses and lichens, in the deep furrows
of the shaggy dark brown bark of the tree. Carefully I
plucked it out. Examination with a lens proved that it
was indeed a very beautiful little orchid less than one
half an inch high with yellowish flowers, of a genus un-
known to me. Calling to Francisco to stop his cooking,
I put the little plant in my note-book and let it fall to
him. All other thoughts dismissed, we spent the rest of
the afternoon searching the trunks of all the trees in the
vicinity, but only a few additional specimens of the or-
chid were found.
[ 54 |
‘It is a coincidence that here occurred examples of
the two extremes of the jungle plants which interest me
most—the most diminutive of epiphytic orchids and the
most gigantic of trees growing together, my interest in
the latter leading to the discovery of the former.’
Brazi.: Upper Rio Negro drainage-area, Rio Dimiti, at the base
of Mt. Dimiti, epiphytic in cracks of the bark of Cunuria crassipes,
flowers yellow, May 12-19, 1948, Richard Evans Schultes & Francisco
Lopez 10003 (Tyrer in Herb. Ames No. 65206),
Epidendrum flexuosissimum C. Schweinfurth
sp. nov.
Herba parva, epiphytica, crassior. Rhizoma repens.
Caules breves, robusti, approximati, vaginis tubulatis
vel vaginarum fibris omnino tecti, apice bifoliati. Folia
subopposita, late patentia, ovalia vel orbicularia, sessilia.
Inflorescentia supra paniculata, laxe multiflora, cum pe-
dunculo rachideque valde fractiflexa. Flores parvi, cum
segmentis late patentibus. Sepalum dorsale elliptico-
lanceolatum vel oblongo-lanceolatum, acuminatum. Se-
pala lateralia paulo majora, obliquissime elliptico-lan-
ceolata, complicato-acuminata. Petala linearia, leviter
sigmoidea, dense ciliata. Labellum columnae valde ad-
natum; lamina carnosa, medio trilobata; lobi laterales
erecto-incurvi, aliformes; lobus medius_ triangularis,
porrectus,
Plant small, rather stout, about 8-17 em. high. Rhi-
zome creeping (usually fragmentary in our specimens).
Roots numerous, fibrous, glabrous, slender, commonly
unbranched. Stems short, stout, suberect to lightly flex-
uous, approximate and spreading from the rhizome, 1-
or 2-jointed, entirely concealed by two or three tubular
imbricating sheaths or the fibres of sheaths, about 1-3
em. long, 2-leaved at the apex. Leaves subopposite,
widely spreading, oval to orbicular, sessile, apparently
always rounded at the apex and sometimes with a small
[ 55
mucro, coriaceous, about 1.5-8.6 cm. long, up to 2.5
cm. wide. Inflorescence divaricately paniculate, much
surpassing the leaves, with the peduncle and rachis mark-
edly fractiflex; peduncle 3-4.5 cm. long, complanate
and narrowly bialate with two to four prominent spread-
ing bracts which are conduplicate, broadly winged on
the back and 1.2 em. or less long. Bracts subtending
the branches of the panicle similar but smaller. Panicle
loosely several- to many-flowered, with spreading or re-
curved few-flowered branches. Flowers small, pale green-
ish yellow, with spreading segments. Dorsal sepal elliptic-
lanceolate or oblong-lanceolate, acuminate or short-
acuminate, 3-nerved, about 8 mm. long and 2.2—2.9 mm.
wide, dorsally keeled toward the apex. Lateral sepals
very obliquely elliptic-lanceolate, complicate-acuminate
above, concave, 3-nerved, prominently keeled toward
the apex, 8.1-8.4 mm. long measured along the posterior
margin, about 38 mm. wide. Petals linear, lightly sig-
moid, acuminate, 1-nerved, with the margins densely
ciliate, about 7 mm. long and 1.1 mm. wide. Lip adnate
to the column nearly to the apex; lamina fleshy, concave
below, 3-lobed near the middle, cordate at the base, acute
or acuminate at the tip, 4-5 mm. long in the center;
lateral lobes erect-incurved, semiovate or aliform, with or
without a free apex; mid-lobe about equally large with
the lateral lobes, porrect, triangular or ovate-triangular.
Column stout and dilated upward when seen from the
side, subentire at the apex, about 4mm. long at the back.
This little species does not appear to have any near
allies.
Panama: Coclé, Cerro Pajita, hills north of El Valle, at 1100
meters altitude, October 27, 1946, Paul H. Allen 3780 (Tyrer in Herb.
Ames No. 65445).
Epidendrum pajitense C. Schweinfurth sp. nov.
Herba elata. Caulis suberectus, gracilior, vaginis arctis
[ 56 |
tubulatis pustulatis omnino velatus. Folia disticha, late
patentia, superne solum praesentia, oblongo-lanceolata
vel anguste elliptico-lanceolata. Inflorescentiae laterales
et terminales, laxe pluriflorae. Florum segmenta valde
patentia. Sepala elliptico-oblonga, lateralia obliqua. Pe-
tala cuneato-spathulata, subacuta. Labellum columnae
valde adnatum; lamina trifida, cum lobis lateralibus ob-
lique obovato-oblongis et lobo medio flabellato-cuneato,
profunde bilobato. Columna apice quadrialata.
Plant rather tall, “‘saprophytic.’” Rhizome apparently
abbreviated. Roots fibrous, numerous, rather stout, gla-
brous. Stem suberect, rather slender, with one strict
flower-bearing branch above (the main stem broken off),
entirely concealed by close tubular pustulose sheaths
which are leaf-bearing above, about 49 cm. high. Leaves
several to numerous, distichous, spreading, oblong-lan-
ceolate or narrowly elliptic-lanceolate, obtuse to sub-
acute, sessile at the shortly cuneate base, 8.5-6.2 em.
long, about 1.3 cm. or less wide, coriaceous. Inflores-
cences lateral and terminal; the lateral on the upper part
of the stem, racemose, short, loosely few-flowered; the
terminal one loosely paniculate with ascending few-
flowered branches. Floral bracts minute, concave, ovate
or oblong-ovate. Flowers medium-sized, rose-pink, with
widely spreading more or less convex segments. Dorsal
sepal oblong or elliptic-oblong, broadly obtuse to sub-
acute, about 7-nerved, rather fleshy, about 9.3 mm. long
and 3.9 mm. wide. Lateral sepals similar, obliquely
elliptic-oblong, obtuse to subacute with a blunt dorsal
mucro, about 10.5 mm. long, measured from the base of
the dorsal margin to the tip, and 4 mm. wide, about 7-
nerved. Petals cuneate-spatulate, abruptly rounded
above with a subacute apex, 3-nerved with the lateral
nerves branching, slightly oblique, about 10 mm. long
and 3.2 mm. wide above. Lip adnate to the column up
[ 57 ]
to the tip; lamina 3-parted, much exceeding the sepals,
submembranaceous; lateral lobes widely spreading, ob-
liquely obovate-oblong, with a more or less truncate and
obscurely lobulate outer margin, separated by a pair of
small complanate calli, about 7.5 mm. long through the
middle lengthwise and 3.9 mm. wide above; mid-lobe
porrect, narrowly flabellate-cuneate, conspicuously bi-
lobed with a minute blunt apicule between the oblong-
rounded lobules and with a small callus at the base be-
tween the lateral calli, about 8.2 mm. long to the tip of
an apical lobule and 7.2 mm. wide across the apical lob-
ules. Column short, very stout and dilated upward when
viewed from the side, about 6.5 mm. long at the back
including the erect 4-lobed wing, with the middle lobes
oblong-subquadrate and irregularly lobulate on the trun-
cate apex.
This species has three rather close allies. Hpidendrum
exasperatum Reichb.f. differs in having smooth cauline
sheaths, verrucose outer surfaces of the sepals and flowers
of another color. 17. Schumannianum Schltr. has broader
leaves, broader petals, relatively smaller lateral lobes of
the lip and spotted flowers. 2. verrucosum Sw. var.
myrianthum (Lindl.) Ames & Correll shows much nar-
rower and relatively longer leaves and a lip with dissim-
ilar details.
Panama: Coclé, Cerro Pajita, hills north of El Valle, at 1100 me-
ters altitude, October 27, 1946, ‘“saprophytic,’’ in dense shade, Paul
H. Allen 3784 (Tyrer in Herb. Ames No. 65444).
Epidendrum strictiforme C. Schweinfurth sp. nov.
Herba mediocris, epiphytica, recta. Rhizoma abest.
Caulis in siccitate crassus, foliorum vaginis ancipitibus
omnino vestitus. Folia disticha, quinque ut videtur, ovata
vel oblongo-ovata, apice rotundata, basi lata amplexi-
caulia. Inflorescentia erecta; pedunculus spathis condu-
plicatis, erectis, imbricatis, duabus vel tribus omnino
[ 58 |
obtectus; racemus dense multiflorus. Flores parvi. Se-
palum dorsale oblanceolatum, acutum. Sepala lateralia
oblique oblongo-oblanceolata, acuta vel breviter acumi-
nata. Petala oblanceolato-linearia, leviter incurva, acuta
vel breviter acuminata. Labellum columnae valde ad-
natum; lamina trilobata, carnosa; lobi laterales rotun-
dato-dolabriformes, extus irregulariter crenulati; lobus
medius ovatus, acutus. Columna generis, recta.
Plant medium-sized for the genus, stout, epiphytic,
up to 87 cm. tall. Roots and rhizome not present. Stem
apparently stout, entirely concealed by strongly compla-
nate tubular leaf-sheaths. Leaves five, loosely disti-
chous, ovate or oblong-ovate, rounded and minutely bi-
lobed at the apex, amplexicaul at the broad base, about
5-7.5 em. long, 2.8-3.4 cm. wide, spreading. Inflores-
cence erect, racemose above; peduncle apparently about
11 em. long, entirely concealed by two or three strictly
erect, conduplicate, imbricating spathes which are
rounded to acute above ; raceme densely many-flowered ;
floral bracts lax, linear-lanceolate, the lower ones about
equaling the slender pedicellate ovary. Flowers small,
pale brownish green flushed with purple, spreading.
Dorsal sepal oblanceolate or elliptic-oblanceolate, acute,
5-nerved, about 9-10 mm. long and 3-3.4 mm. wide
above. Lateral sepals obliquely oblong-oblanceolate,
lightly sigmoid, acute or short-acuminate, longitudinally
concave, 5-nerved, about 10.3-11 mm. long on the pos-
terior margin and 38-3.3 mm. wide just above the middle.
Petals oblanceolate-linear, lightly incurved, acute or
short-acuminate, l-nerved, about as long as the dorsal
sepal and 1 mm. wide above. Lip adnate to the column
up to its apex; lamina sharply 3-lobed, fleshy, cordate
at the base, about 5-5.8 mm. long in the center and
6.4-7.7 mm. wide across the lateral lobes; lateral lobes
rounded-dolabriform, with irregularly crenate outer mar-
[ 59 |
gins; mid-lobe ovate, acute, protuberant, very fleshy,
about 3-3.5 mm. long and 2.5—2.9 mm. wide at the base ;
dise with a pair of small fleshy conical calli at the base.
Column straight, dilated above in front, retuse in the
center at the summit, about 6.5-7 mm. long on the dor-
sal surface, produced on each side into an obliquely
semiorbicular-ovate lobe.
This species apparently lacks any close allies. It may
be related to Mpidendrum sarcodes Lindl., but differs in
having the peduncle entirely concealed by the spathes,
and in having acute sepals, ete.
Peru: Hudnuco, Carpish, at 2800 meters altitude, on a tree in
cloud forest, leaf not thick but very hard and stiff, tip of column
white, September 1946, F. Woytkowski 37014 (Tyre in Herb. Ames
No. 65451).
Brassavola ovaliformis C. Schweinfurth sp. nov.
Herba epiphytica, pro genere mediocris, deserticola.
Rhizoma repens, nodulosum. Caules approximati, plus-
minusve elongati, vaginis tubulatis imbricatis scariosis
omnino obtecti. Folia subteretia, gracilia, elongata, in
siccitate arcuata. Flores duo, grandes, in pedunculi apice
approximati. Sepala petalaque late patentia, valde sim-
ilia, lanceolato-linearia, sensim angustata, apice acuta.
Labellum multo brevius, inferne leviter involutum, ex-
pansum ovale-ovatum, acutum, ecarinatum, margine in-
tegro. Columna perbrevis, apice trialata, cum alis later-
alibus faleato-lanceolatis et recurvis.
Plant epiphytic. Roots fibrous, covered with a stout
glabrous velamen. Rhizome creeping, nodulose, consist-
ing of the approximate swollen bases of the stems. Stems
crowded, up to 12.7 cm. or more long (the longest one
incomplete), several-jointed, entirely enveloped by long
tubular scarious imbricating sheaths, gradually dilated
upward, 1-leaved at the apex. Leaves very slender, sub-
terete, channelled, arcuate in the dried specimen, about
[ 60 ]
27.2-88.2 cm. or more long (the longest blade incom-
plete), apparently about 3 mm. in diameter. Peduncle
exceeding 2.5 cm. in length (incomplete), with two flow-
ers at the apex. Floral bracts very small, lanceolate-
ovate, acuminate, deeply concave, amplexicaul, much
shorter than the pedicellate ovary which is about 4 cm.
long. Flowers large with widely spreading segments.
Dorsal sepal lanceolate-linear, long-narrowed to an acute
tip, about 5.8-5.8 em. long and 5.5 mm. wide near the
base. Lateral sepals closely similar, about 5.9 ecm. long
and 5 mm. wide below. Petals similar to the sepals,
linear-lanceolate, about 5—-5.8 em. long and 5.4 mm.
wide below. Lip much shorter than the other segments,
oval-ovate or lanceolate-oval in outline when expanded,
about 4.7-4.9 em. long and 2.9 em. wide across the mid-
dle, acute, ecarinate, with the lower half gradually in-
rolled in natural position, the margins being entire.
Column minute, with an erect 8-lobed wing at the apex,
about 6.5 mm. high at the back; lateral wings falcate-
lanceolate and recurved ; middle wing low and irregular,
abruptly extended in the center into a ligulate, apically
truncate and denticulate projection; anther oblong-
ellipsoid, 2-celled, with four narrowly oblong-ellipsoid
pollinia in each cell.
This species appears to have no near allies. In the
whiteness of the flowers and in the apex of the column
it resembles the widespread Brassavola nodosa (.)
Lindl., but it lacks the elongated narrow serrated basal
portion of the lip of that species. B. Perriniit Lindl.,
from Brazil, differs in having smaller green flowers, a
somewhat carinate lip and dissimilar column-wings.
Peru: Amazonas, La Peca, northeast of Jaen, at 1000 meters al-
titude, on dwarf trees in dry “‘desert forest full of cacti,’’ flower uni-
formly white, the lip having a median greenish yellow line above and
ee, e eas
beneath, interior parts ... pale green save the whitish column and
the two formations immediately below, which are whitish with a brown
[ 61 ]
hue, leaves dark green, tapering, stiff, round like thick wire,’’? No-
vember 18, 1947, F. Woytkowski 37016 (Tyre in Herb. Univ. Calif.).
Odontoglossum angustatum Lindley in Bot.
Reg. 23 (1887) sub t. 1992; Fol. Orch. Odontoglossum
(1852) 17, no. 48, non O. angustatum Lindley Orch.
Linden. (1846) 17, no. 90; Bateman Monog. Odontog.
(1874) t. 26.
Odontoglossum tetraplasium Reichenbach filius in
Gard. Chron. n.s. 8 (1875) 558.
Odontoglossum bellum Schlechter in Fedde Repert.
Beih. 9 (1921) 108; ex Mansfeld in Fedde Repert.
Beih. 57 (1929) t. 127, nr. 499.
Odontoglossum Loesenerianum Schlechter in Fedde
Repert. Beih. 9 (1921) 110; ex Mansfeld in Fedde
Repert. Beih. 57 (1929) t. 128, nr. 502.
A sketch of the panicle with a drawing of a solitary
flower (natural size) and a single enlarged lip of Odonto-
glossum tetraplasium from the Reichenbach Herbarium
shows that this concept cannot reasonably be separated
from O. angustatum Lindl. (1837), as represented by a
photograph of the type specimen from the Lindley Her-
barium bearing a pen drawing of the callus on the lip.
The vegetative parts were lacking in the type specimens
of both species, but these were supplied for O. angusta-
tum by a subsequent collection mounted with the Lind-
ley type.
Judging from the description and floral analysis, Odon-
toglossum bellum is also a form of this variable species.
O. bellum appears to be quite as large throughout as
typical O. angustatum, but the sepals are not so elongate-
acuminate as in many forms of the latter. The lip which
is described and shown as obtuse and apiculate, is not very
dissimilar to the acute lip of Vargas 2888 and Vargas
3664 which have been determined as O. angustatum.
[ 62 |
The floral analysis of Odontoglossum Loesenerianum
indicates that this concept is surely referable to O. an-
gustatum as first described by Lindley, the only apparent
discrepancies being that the bracts are noted as being
equal to or a little exceeding the ovary (they are about
half as long in O. angustatum) and the flowers are de-
scribed as somewhat smaller. This difference in size is
probably due to an immature condition.
In the several collections here referred to O. angusta-
tum, there appears to be a wide variation both in vege-
tative and floral size and in the form of the lip. The most
striking variations from the usual form are seen in Var-
gas 2888, where the obviously immature flowers are
smaller than those of O. Loesenerianum and have a lip
which is more or less lightly pandurate; and in Weber-
bauer 7797, where the carinate calli at the base of the
lip are broader, blunter and thicker than in the usual
form. All of the Peruvian collections included in this
species, however, have a general similarity of lip calli,
but there is apparently lacking from all of them one extra
pair of tubercles present in the type.
It seems to me, therefore, that Odontoglossum angus-
tatum is a conspicuous example of the polymorphism
that makes the tropical orchids so difficult.
Odontoglossum aureo-purpureum (as auropur-
pureum) Reichenbach filius in Linnaea 22 (1849) 848;
Lindley Fol. Orch. Odontoglossum (1852) 15, no. 44.
Odontoglossum compactum Reichenbach filius in Gard.
Chron. n.s. 3 (1875) 492.
Odontoglossum Koehleri Schlechter in Fedde Repert.
Beih. 9 (1921) 109; ex Mansfeld in Fedde Repert.
Beih. 57 (1929) t. 128, nr. 501.
The concept Odontoglossum compactum is based in part
on Peruvian specimens collected by W. Lobb which
[ 63 ]
were formerly referred by Lindley (Fol. Orch. Odonto-
glossum p. 15) to O. aureo-purpureum. It was separated
from the earlier species by its more compact densely
flowered panicles of larger flowers. However, two recent
Peruvian collections (Vargas 2879 and Metcalf 30742)
have the loose panicles seen in the typical Venezuelan
O. aureo-purpureum, but they have larger flowers than
those of the type which they appear to match in floral
details.
Odontoglossum Koehleriis described as having a rather
densely flowered panicle, like O. compactum; but its
floral segments, except for being slightly broader, coin-
cide well with those of typical O. aureo-purpureum.
It seems to me, therefore, that O. awreo-purpureum
is a widely variable species which includes the above
concepts.
Odontoglossum brevifolium Lind/. var. Weber-
bauerianum (Kriinzl.) C. Schweinfurth comb. nov.
Oncidium Weberbauerianum Krinzlin in Engler Bot.
Jahrb. 37 (1906) 389.
Cyrtochilum Weberbauerianum Krinzlin in Notizbl.
Bot. Gart. Berlin 7 (1917) 95, nomen nudum, in clavi;
in Engler Pflanzenreich LV. 50, pt. 2 (Heft 80) (1922)
60, fig. 5H, a-c.
Odontoglossum Weberbauerianum Schlechter in Fedde
Repert. Beih. 27 (1924) 109.
In Fedde Repert. Beih. 9 (1921) 170, Dr. Schlechter
cites Oncidium Weberbauerianum as a synonym of the
well-known Odontoglossum brevifolium Lindl.
I have not seen any figure of Oncidium Weberbaueri-
anum, but, after examining an iso-type collection of
Odontoglossum brevifolium, there appear to be several
marked differences between that species and the descrip-
tion of Cyrtochilum Weberbauerianum (supplemented by
[ 64 |
a floral analysis). It seems to me, therefore, that the
above varietal designation is preferable, for the reasons
indicated in the following table of comparisons.
Odontoglossum brevifolium
Pseubobulbs ovoid to cylindric-
ovoid, up to 5 em. long.
Leaves oval to oblong-elliptic,
up to 15 em. long and 7.6 cm.
wide.
Sepals suborbicular to round-
obovate, little longer than broad.
Lip deeply bilobed at the apex.
Oncidium Weberbauerianum
Pseudobulbs “‘linear,’’ up to 10
em. long.
Leaves oblong, up to 20 em. long
and 4.5 em. wide.
Sepals oblong, much longer than
broad.
Lip obscurely lobulate in front.
Odontoglossum flavescens fPo/fe in Orch. Rev.
12 (1904) 92.
This species was too inadequately described to furnish
a definite concept that can be visualized or keyed. The
description, which gives no hint of vegetative parts,
merely states that the flowers are clear yellow, that the
segments are elliptical-oblong and subconnivent, while
the blade of the lip is suberect, rounded below, with an
oblong obtuse apex and has a very large 2-lobed callus
appressed to the base of the column. It is noted as being
allied to O. retuswm Lindl., and was introduced by
Messrs. F. Sander & Co., presumably from Ecuador
and Peru.
A photograph of the type specimen in the Herbarium
at Kew shows fragments of an elliptical-oblong pseudo-
bulb which is about 8.5 em. long and appears to be bifo-
liate or trifoliate. The single leaf shown is oblong-linear,
short-acuminate, slightly narrowed to a sessile base, over
51cm. long and about 2.4cm. wide, with the mid-nerve
very prominent beneath. The inflorescence shows a very
loosely branched panicle with two horizontally spreading
branches. ‘The flowers are distant, secund, on long ped-
icels (up to 2 cm. long) much exceeding the minute
bracts, and the segments appear to be about 1 cm. long.
[ 65 ]
Even with this additional information, however, it
seems advisable to consider this concept among the ob-
scure species.
Odontoglossum mystacinum (Lindl.) Lindley
Fol. Orch. Odontoglossum (1852) 6, no. 14.
Cyrtochilum mystacinum Lindley in Bot. Reg. 24
(1838) Mise. 30, no. 38; in Bot. Reg. 25 (1839) t. 62.
Odontoglossum rigidum Lindley in Benth. Pl. Hartw.
(1844) 152; Fol. Orch. Odontoglossum (1852) 7, no.
Ve
Judging by an excellent photograph of the type of
Odontoglossum rigidum from the Lindley Herbarium,
this concept cannot reasonably be separated from the
species depicted as Cyrtochilum mystacinum in the Botan-
ical Register 25, t. 62. A slight difference is that the
pseudobulb of the latter concept is represented as uni-
foliate, whereas that of O. rigidum is bifoliate with very
unequal leaves. Recent Peruvian collections which are
referable to this concept have sometimes unifoliate and
sometimes bifoliate pseudobulbs. Although O. rigidum
has a more branched and fractiflex panicle than that of
Cyrtochilum mystacinum, the plate was obviously drawn
from an immature specimen. A minor difference is that
the wings of the column in the latter concept are rather
broad and multifid, whereas they are drawn as rather
narrow and serrate in O. rigidum.
Both species, however, have similar pseudobulbs and
yellow flowers with almost identical segments. The se-
pals are lanceolate, the petals ovate-lanceolate and the
lip pandurate-obovate. Moreover, the column wings are
not easily seen in the specimens examined and appear to
be variable.
This species appears to be limited to Peru, Ecuador,
and perhaps Bolivia.
[ 66 |
Odontoglossum obscurum C. Schweinfurth nom.
nov.
Mesospinidium Wallis Reichenbach filius in Linnaea
41 (1876) 106, non Odontoglossum Wallisii Linden &
Reichb. f. (1870).
As illustrated by a series of floral analyses from the
Reichenbach Herbarium, Mesospinidium Wallisii has the
basal part of the lip parallel to the column with the an-
terior half abruptly reflexed. It seems, therefore, to be
clearly referable to the genus Odontoglossum.
Since the specific epithet, Wadllisii, has already been
used under Odontoglossum, the new name obscurum is
proposed, in allusion to the fact that the pseudobulb and
leaves were unknown when the species was described.
Except for the lip, which is described and illustrated
as retuse at the apex and emarginate in the middle [of
each side], this concept might reasonably be referred to
Odontoglossum longifolium Lindl.
Odontoglossum Wyattianum G. Wilson in Orch.
Rev. 36 (1928) 47.
In the diagnosis of this species there is no description
of the vegetative parts of the plant nor of the inflores-
cence. With regard to the flowers, only the size and color
are noted; there is no mention of the exact shape of the
parts. Finally, the precise origin of the species is doubt-
ful, for the description merely states that ‘‘the plant had
been obtained from the Rev. Paul Wyatt, Bedford, to
whom it had been forwarded by a friend in Peru, doubt-
less its native country. ”’
It seems advisable, at the present time, to relegate
this concept to the class of obscure species.
Oncidium falcipetalum Lindley Orch. Linden.
(1846) 14, no. 76; Fol. Orch. Oncidium (1855) 5, no. 7:
[ 67 ]
Cogniaux Dict. Icon. des Orch. Oncidium (1899) t. 21.
Oncidium Pavoni Reichenbach filius ex Lindley Fol.
Orch. Oncidium (1855) 5, sub no. 7, nomen nudum;
Kriinzlin in Engler Pflanzenr. LV. 50, pt. 2 (Heft 80)
(1922) 38.
Oneidium ionodon Reichenbach filius in Linnaea 41
(1876) 23.
Oncidium Davisn Reichenbach filius in Linnaea 41
(1876) 24.
Cyrtochilum faleipetalum Krinzlin in Engler Pflanzenr.
IV. 50, pt. 2 (Heft 80) (1922) 87, fig. 1 G, a-e.
Cyrtochilum Pavoni Krinzlin in Engler Pflanzenr.
IV. 50, pt. 2 (Heft 80) (1922) 388.
Oneidium Pavoni, although lacking any original diag-
nosis, has been considered by the several writers men-
tioning the name as a form of O. falcipetalum.
Judging from the description (limited to the flower),
the Peruvian Oncidium ionodon differs from O. falcipet-
alum only in having an emarginate dorsal sepal, clawed
petals and porrect violet lateral lobes of the lip. How-
ever, the plate of O. falcipetalum illustrated in Cogniaux’
work (I.c.) shows a crisped dorsal sepal, which might well
be taken for emarginate, and petals that appear to be
more or less clawed. Also, in specimens reasonably re-
ferred to O. falcipetalum, the petals (when examined
closely) are very shortly clawed. In general, too, the
color of the flower attributed to O. tonodon coincides
with that shown in the plate indicated.
Onecidium Davis, of which there is in the Ames Her-
barium a tracing of the floral analysis from the Reichen-
bach Herbarium, is almost an exact counterpart of O.
ionodon, except that the dorsal sepal is noted as acute,
the petals as very shortly clawed, and the lateral lobes
of the lip deflexed.
It is a strange coincidence that both O. ionodon and
[ 68 |
O. Davisii are cited as synonyms of one species, Cyrto-
chilum monachicum, by Krinzlinin Engler Pflanzenr. IV.
50, pt. 2 (Heft 80) (1922) 47. However, the latter con-
cept, based upon Oncidium monachicum Reichb.f., is de-
scribed as having the dorsal sepal reniform, and thus is
quite dissimilar to that of O. falcipetalum or its forms.
Apparently Oncidium falcipetalum is a variable species
as regards floral size and the contour of the petals which
varies from flat to conduplicate.
Originating in Venezuela (Merida), this species is also
recorded from Colombia and Peru.
Oncidium heteranthum Poeppig & Endlicher Nov.
Gen. ac Sp. 1 (1886) 34, t. 60; Cogniaux in Martius FI.
Bras. 8, pt. 6 (1905) 878, t. 87; Krinzlin in Engler
Pflanzenr. IV. 50, pt. 2 (Heft 80) (1922) 175.
Oneidium bryolophotum Reichenbach filius in Gard.
Chron. (1871) 788; Krinzlin in Engler Pflanzenr. LV.
50, pt. 2 (Heft 80) (1922) 181.
Oncidium inops Cogniaux & Rolfe in Journ. des Orch.
3 (1893) 846; 4 (1893) 74.
Oneidium megalous Schlechter in Fedde Repert. 9
(1911) 30.
The above species is extremely variable. This fact has
led to the proposal of several later concepts, none of
which, however, seems valid.
Oneidium bryolophotum, a native of Costa Rica and
Panama, and represented in the Ames Herbarium by
Reichenbach’s detailed analyses, by authentic material
examined by Kriinzlin, and by many Central American
collections, cannot logically be separated from the earlier
O. heteranthum trom Colombia, Peru and Bolivia.
In general, the Central American plants referred to
O. bryolophotum have a mid-lobe of the lip which is rel-
atively larger than that of O. heteranthum, but one Costa
[ 69 |
Rican collection (Standley 33741) has a lip which is quite
similar to that of the South American O. heteranthum.
One Peruvian collection of O. heteranthum (Vargas
2531) varies from the usual form in lacking the separate
calli on the lateral (or basal) lobes of the lip and in hav-
ing the apex of the column-wings narrowed to an acu-
minate point rather than having the typical more or less
broad apex. Another Peruvian collection (Schunke s.n.,
Herb. Field Mus. No. 571665) has a row of detached
calli extending onto the lateral lobes on each side of the
middle callus.
Other differences noted in the various collections ap-
pear in the vegetative size, in the oblong-ovoid to cylin-
dric pseudobulbs, and in the leaves which vary from
linear-oblong to elliptic.
The other concepts cited above have already been re-
duced to the synonymy of O. bryolophotum.
Oncidium incarum (Ariinzl.) C. Schweinfurth
comb, nov.
Cyrtochilum Incarum Krinzlin in Engler Pflanzenr.
IV. 50, pt. 2 (Heft 80) (1922) 58.
Since it seems unwise to regard the concept Cyrtochi-
lum as distinct from the large and variable genus Oncid-
ium, the above transfer is necessary.
Oncidium macranthum Lindley var. hastiferum
(Reichb.f. & Warse.) C. Schweinfurth comb. nov.
Oncidium hastiferum Reichenbach filius & Warscewicz
in Bonpl. 2 (1854) 102; Reichenbach filius in Walp.
Ann. 6 (1863) 708.
Cyrtochilum hastiferum Krinzlin in Engler Pflanzenr.
IV. 50, pt. 2 (Heft 80) (1922) 34.
Judging by the descriptions of Oncidium hastiferum
and by a drawing in the Reichenbach Herbarium, made
from cultivated material, it appears preferable to con-
[ 70 ]
sider this concept as a variant of the widespread O.
macranthum. It seems to differ from O.macranthum only
in having one large median keel flanked by a smaller
lamella on each side (instead of three equal lamellae) and
in having narrower ligulate wings on the column.
Oncidium obryzatum Reichenbach filius & Warsce-
wicz in Bonpl. 2 (1854) 108; Regel in Gartenfl. 27 (1878)
t. 925; Kriinzlinin Engler Pflanzenr. IV. 50, pt. 2 (Heft
80) (1922) 239, fig. 20 B, a-d.
The Peruvian collection cited below shows several dis-
crepancies from typical examples of O. obryzatum in the
Ames Herbarium.
The pseudobulb is bifoliate, not unifoliate as described.
The leaves are longer than exemplified, being up to 52.5
em. long. The branches of the peduncle appear to be
rather more distant than otherwise shown, being about
5-6.5 cm. (instead of 2-4 cm.) apart. The sepals and
petals, although closely similar to those of the typical
form, are somewhat abruptly acute, rather than obtuse
or rounded or even retuse. The narrow middle part of
the lip is somewhat broader than usual and the anterior
portion or middle lobe is narrower relative to the basal
portion. The column wings are merely acute, not acu-
minate, above.
In view of the general agreement of this specimen
with the typical form, however, this collection is incor-
porated with O. obryzatum.
Peru: Cajamarca, Cutervo, about Socota, on Socota River, at 2800
meters altitude, epiphyte, December 11, 1938, H. E. Stork & O. B.
Horton 101852.
Oncidium pyramidale Lindley in Ann. & Mag.
Nat. Hist. 15 (1845) 384; Fol. Orch. Oncidium (1855)
29, no. 98; Veitch Man. Orch. Pl., pt. 8 (1892) Oncid-
ium 73; Kranzlin in Engler Pflanzenr. IV. 50, pt. 2
(Heft 80) (1922) 195, fig. 16 J, a-c.
[71 ]
Oncidium chrysopyramis Reichenbach filius & Warsce-
wicz in Bonpl. 2 (1854) 108; Lindley Fol. Orch. On-
cidium (1855) 29, no. 97; Veitch Man. Orch. PIl., pt.
8 (1892) Oncidium 25; Krinzlin in Engler Pflanzenr.
LV. 50, pt. 2 (Heft 80) (1922) 196, fig. 16 F, a-d.
Judging from drawings of the habit and floral analyses
of Oncidium chrysopyramis from the Reichenbach Her-
barium, supplemented by the descriptions, it does not
seem reasonable to separate this concept from the earlier
O. pyramidale. The latter species appears to differ from
O. chrysopyramis in having a longer panicle with the
lower branches often compound and many-flowered, not
simple and 8- to 5-flowered—a character which is varia-
ble and therefore unimportant as a basis for specific sep-
aration.
The basal portion of the lip is described as being
broader than the anterior portion in both species, con-
trary to remarks made by Reichenbach following his
description of O. chrysopyramis.
Again, the wings of the column, a character which
was considered of great consequence in separating the
concepts, appear to be highly variable in Reichenbach’s
own figures. Indeed, the column-wings shown in the
pen drawing on the type collection of O. pyramidale
appear to be a close approximation to some of those de-
picted by Reichenbach for O. chrysopyramis.
Oncidium superbiens Reichenbach filius in Lin-
naea 22 (1849) 843; Hooker filius in Bot. Mag. 98 (1872)
t. 5980; Warner & Williams Orch. Alb. 6 (1887) t. 276.
Oncidium aemulum Reichenbach filius & Woarscewicz
in Bonpl. 2 (1854) 102.
Oncidium inferlobum hort. ex Gard. Chron. (1872)
904, in synon.
Oncidium undulatum Warner & Williams Orch. Alb.
[ 72 ]
8 (1889) t. 368, non O. undulatum (HBK.) Lindl.
(1842).
Cyrtochilum aemulum Wrinzlin in Notizbl. Bot. Gart.
Berlin 7 (1917) 98, nomen nudum, in clavi; in Engler
Pflanzenr. IV. 50, pt. 2 (Heft 80) (1922) 47, fig. 3 C,
a—e.
Cyrtochilum superbiens Krinzlin in Notizbl. Bot. Gart.
Berlin 7 (1917) 93, nomen nudum, in clavi; in Engler
Pflanzenr. IV. 50, pt. 2 (Heft 80) (1922) 49, fig. 3 E,
oe
After a careful comparison between Oncidium super-
hiens (represented by a photograph of the type and the
type description) and O. aemulum (represented by a
photograph of Lindley’s concept, supplemented by the
type description), I am unable to separate the two species.
All the specimens of O. superbiens noted appear to
have the petals yellowish or whitish, more or less barred
below with brown or purplish brown, whereas there is
no record of such coloring in O. aemulum.
The figures of these concepts in the Pflanzenreich
(under Cyrtochilum) confirm my conclusions that they
are conspecific.
Oncidium ventilabrum Reichenbach filius & War-
scewicz in Bonpl. 2 (1854) 101; Lindley Fol. Orch. On-
cidium (1855) 6, no. 12.
Cyrtochilum undulatum Humboldt, Bonpland & Kunth
Nov. Gen. et Sp. Pl. 1 (1816) 849, t. 84; Krinzlin in
Engler Pflanzenr. IV. 50, pt. 2 (Heft 80) (1922) 57.
Oncidium undulatum Lindley Sert. Orch. (1842) sub
t. 48, no. 1; Reichenbach filius & Woarscewiez in
Bonpl. 2 (1854) 108; Lindley Fol. Orch. Oncidium
(1855) 7, no. 15, nee Oncidium undulatum Sims (1804)
nec O. undulatum Salisb. (1812).
Cyrtochilum ventilabrum Kriinzlin in Notizbl. Bot.
[ 73 ]
Gart. Berlin 7 (1917) 92, nomen nudum, in clavi; in
Engler Pflanzenr. IV. 50, pt. 2 (Heft 80) (1922) 39,
fig. 2C, a-c.
In his original description of Oncidium ventilabrum
Reichenbach admitted that this species was close to O.
undulatum Lindl., but he adduced several supposedly
weighty reasons for separating them. Among other fea-
tures he said, ‘‘Zuniichst hat unsre Art zweimal so grosse
Bliithen, die seitlichen Sepala breiter, das obere linger.
Der mit dem Fuss der Siitule verwachsene Lippennagel
is viel linger... .”°
In the Ames Herbarium there are several records of
Oncidium ventilabrum (a drawing of a flowering branch
and numerous floral analyses) from the Reichenbach
Herbarium, and these show a rather striking agreement
both in size and form of sepals and petals with those
shown in the plate of the type of Cyrtochilum undulatum
(l.c.). Moreover, the lip is depicted as sessile in both
concepts. The form of the lip and the calli at the base
of the lamina are supposed to be different in the two
species, but in the drawings of O. ventilabrum there is a
rather wide range in these features. Furthermore, in the
remarks about O. undulatum (in Bonpl. 2, p. 108) Reich-
enbach described a lip-callus which is rather similar to
that shown in O. ventilabrum.
It appears to be the wise procedure to eliminate the
necessity of straining for supposed differences between
these two concepts and to consider them conspecific.
However, since there is an earlier and different On-
cidium undulatum, it is necessary to reject this specific
epithet and to adopt the next earlier name, Oncidiwm
ventilabrum.
Sigmatostalix Reichenbach filius in Bot. Zeit. 10
(1852) 769; Schlechter in Fedde Repert. 15 (1918) 139;
[ 74 ]
Krinzlin in Engler Pflanzenr. LV. 50, pt. 2 (Heft 80)
(1922) 301.
Petalocentrum Schlechter in Fedde Repert. 15 (1918)
144; Kranzlin in Engler Pflanzenr. [V. 50, pt. 2
(Heft 80) (1922) 312.
The concept Petalocentrum was separated from the
variable genus Sigmatostaliv by reason of three alleged
differences (Cf. Schlechter in Fedde Repert. 15 (1918)
145), viz. the sessile lip, the spur-like outgrowth at the
base of the petals and the ascending rostellum.
The characterization of the lip in the type description
of Sigmatostalix (1.c), to be sure, specifies ‘‘labellum un-
guiculatum.’’ However, the lip in the typical species,
S. graminea (as Specklinia graminea Poepp. & Endl. in
Nov. Gen. ac Sp. 1 (1836) 51, t. 89 B), appears to be
cuneate below to a sessile base; and, even if the analysis
is discarded as being inadequate (on the basis of later
representations), that organ seems to be at most only
very shortly unguiculate.
At any rate, some species which were described as be-
longing to Sigmatostalivx, such as the Central American
S. hymenantha Schitr., have the basal part of the lip
more or less cuneate to a sessile base (apparently very
broadly cuneate). And other species, such as S. macro-
bulbon Kriinzl., have an absolutely sessile lip. In the face
of such wide variation, therefore, it seems advisable to
discard this character, at least as to its generic value.
As regards the spur-like outgrowth on the petals, this
character appears to be quite absent in some species with
a strikingly sessile lip which were referred to Sigmato-
stalix, such as iS. macrobulbon Krinzl. Nor does this
character appear in S. hymenantha Schltr. with the
broadly cuneate base to the lip. It would seem, there-
fore, that this outgrowth on the petals, seen in one poly-
morphic species of this alliance, is scarcely of generic
weight when taken alone.
Finally, the factor of an ascending rostellum appears
to be a matter of degree, even if not too recondite and
obscure for ordinary recognition in the dried specimen.
It appears to me that the wise course is to relegate
Petalocentrum to the synonymy of the variable genus
Sigmatostala.
The concept, Petalocentrum angustifolum, the only
one of the genus not previously described as representing
Sigmatostalix, was considered identical with Sigmatosta-
liv pusilla Schltr. (cf. Kriinzlin in Engler Pflanzenr. LV.
50, pt. 2 (Heft 80) (1922) 312), and the generic identity
of the concepts Petalocentrum and Sigmatostalia was
once suspected by Kriinzlin (ef. lc. 318).
Sigmatostalix peruviana /folfe in Kew Bull.
(1910) 371.
Sigmatostalix pusilla Schlechter in Fedde Repert. 10
(1912) 392.
Sigmatostalix bicornuta Rolfe in Kew Bull. (1913) 842.
Petalocentrum pusillum Schlechter in Fedde Repert.
15 (1918) 145; Krinzlin in Engler Pflanzenr. IV. 50,
pt. 2 (Heft 80) (1922) 312; ex Mansfeld in Fedde
Repert. Beih. 58 (1980) t. 59, nr. 285.
Petalocentrum angustifolum Schlechter in Fedde Re-
pert. 15 (1918) 145; ex Mansfeld in Fedde Repert.
Beih. 58 (1930) t. 59, nr. 234.
Petalocentrum bicornutum Schlechter in Fedde Re-
pert. Beih. 9 (1921) 179.
All of the concepts cited above appear to be closely
similar vegetatively, although Sigmatostalia pusilla is
described as having markedly shorter pseudobulbs and
leaves than the others. It is significant that the sepals
and petals of all of the concepts are of almost exactly the
[ 76 |
same form and size, the only discrepancy being in the
degree of acuteness of the apex. In 8. peruviana there
is no mention of a horn at the base of the petals (which
occurs in all of the others), but this species seems surely
identical with S§. bicornuta, even the color of the flowers
being nearly identical. The lip seems to vary somewhat
in size in the various species, but it appears to be of very
similar form throughout. A rather wide diversity is de-
scribed in the callus on the lip in the various concepts,
but it is noteworthy that this feature appears to be often
very variable in a single species of Sigmatostalia.
Since, without actual specimens, it would be extremely
difficult to recognize differences between these species,
the wise course suggests their union.
Ornithocephalus gladiatus Hook. var. peruvi-
anus C. Schweinfurth var. nov.
Planta scapis brevioribus folia non valde excedentibus,
sepalorum mucronibus valde prominentibus, sepalo dor-
sali quam sepalis lateralibus majore, ac praesertim labelli
callo margines basales solum paulo superanti a specie
differt.
Plant small, with the general appearance of Ornitho-
cephalus gladiatus Hook., but differing in having scapes
shorter than or subequaling (not markedly exceeding)
the leaves, in having a very conspicuous mucro on the
sepals, in having the dorsal sepal somewhat larger than
the lateral sepals, and particularly in having a less prom-
inent and spreading basal callus on the lip which but
slightly exceeds the margins.
Peru: Junin, La Merced, at about 610 meters altitude, in mon-
tania, on tree trunk, flowers white with green markings, August 10-
24, 1923, J. Francis Macbride 5480 (Tyre in Herb. Field Mus. No.
536520).
<
ILLUSTRATIONS
EXPLANATION OF THE ILLUSTRATION
PLare XII. Cranicurs crrmLasia C. Schweinfurth.
1, plant, natural size. 2, flower expanded, from
the front, four times natural size. 8, half of the
lip, cut longitudinally, from the side, seven times
natural size.
Drawn by E. W. Smiru
PLATE XII
CRANICHIS
~~
(720,
litlab
~
C2
Ca chweinf.
EXPLANATION OF THE ILLUSTRATION
Pirate XIII. Ocromerta pyemara C, Schweinfurth.
1, plant, eight times natural size. 2, dorsal sepal,
sixteen times natural size. 3, lateral sepal, sixteen
times natural size. 4, petal, sixteen times natural
size. 5, lip expanded, from above, thirty-two times
natural size.
Drawn by EK. W. Suiri
PuatE XIII
OCTOMERIA
PYG MACH
vf
(2
C. Schwe
EXPLANATION OF THE ILLUSTRATION
Pirate XIV. EerpenpruM FLEXUOsIssIMUM C. Schwein-
Jurth, 1, plant, three fourths natural size. 2, flower
from the side, natural position, two and one half
times natural size. 3, lip and column from the side,
natural position, three and one half times natural
size. 4, lamina of lip expanded and apex of col-
umn, from the front, five times natural size. 5, dor-
sal sepal, three and one half times natural size.
Drawn by FE. W. Surru
PLaTE XIV
EPIDENDRUM
flexuosr SS Lmnunme
Cu chweinf-
EXPLANATION OF THE ILLUSTRATION
PLate XV. Eprpenprum pasiTensE C.Schweinfurth.
1, plant, one half natural size. 2, flower partially
expanded, from the front, one and one half times
natural size. 3, lip and column from the side, nat-
ural position, one and one half times natural size.
4, lamina of lip expanded, from the front, twice
natural size. 5, lateral sepal, twice natural size.
Drawn by E. W. Surru
PLaTE XV
me '
A \= :
M, | - EPIDENDRUM
Pes # tCMNSE
pai
CS chweinfr
EXPLANATION OF THE ILLUSTRATION
Pirate XVI. Eprpenprum strictirorMe C,Schwein-
Jurth. 1, plant, seven sixteenths natural size. 2,
flower expanded, from the front, two and one half
times natural size. 3, column and lip from the side,
natural position, two and one half times natural
size. 4, lateral sepal, two and one half times nat-
ural size.
Drawn by E. W. Suitru
PLATE XVI
EPIDENDRUM
strictiforme
C8 chweinf-
EXPLANATION OF THE ILLUSTRATION
Prare XVII. Brassavota ovatirormis C.Schwein-
Jurth. 1, plant, one half natural size. 2, column
from the side, twice natural size. 3, column from
above, twice natural size. 4, lip expanded, from the
front, one half natural size.
Drawn by KE. W. Smiru
PLatTE XVII
BRASSAVOLA
———
ovaliformis
~
fg
C. Schwe
BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY
CampripGr, Massacuusetts, Fesruary 6, 1950 Voi. 14, No. 4
STUDIES IN THE GENUS HEVEA III
BY
RicHarp Evans ScHuutres!
ON THE USE OF THE NAME HEVEA BRASILIENSIS
For some time, it has been believed by certain author-
ities that Hevea brasiliensis, the name long used to de-
note the well known cultivated rubber tree, is untenable
if the International Rules of Botanical Nomenclature
be strictly interpreted.
Several botanists have written on the complicated
problem which underlies this presumed untenability,
treating it from differing points of view and with diverg-
ing conclusions. This divergency of opinion has led to
uncertainty amongst taxonomists not only concerning
the actual status of the name Hevea brasiliensis, but also
as to the authorities to whom it should be attributed.
In 1858, Baillon (Etude Euphorb. (1858) 326) pointed
out that the name Stphonia brasiliensis HBK. had been
applied apparently to two distinct plants: one from the
Orinoco and one from the lower Amazon. He proposed
to reserve Stphonia brasiliensis for the latter—which is
our cultivated species—and published a new name (:S?-
phonia Kunthiana Baill.) for the former.
Later, in 1900, Warburg (Kautschukpflanzen (1900)
'Botanist, Bureau of Plant Industry, Soils, and Agricultural Engi-
neering, Agricultural Research Administration, United States Depart-
ment of Agriculture; Research Fellow, Botanical Museum, Harvard
University.
[ 79 ]
26) discussed this same problem and resolved, in ac-
cordance with what appeared to him to be priority, to
keep the name Hevea brasiliensis for the Orinoco species
and to give the cultivated plant an entirely new name
(Hevea Sicberi Warb.).
Shortly thereafter, Huber (in Bull. Soc. Bot. France
49, ser. 2 (1902) 43) thought it a better policy to con-
serve Hevea brasiliensis for the cultivated species, and
he actually made the proposal that it be so conserved,
In 1905, Ule studied the circumstances and also con-
cluded (in Engler Bot. Jahrb. 85 (1905) 664) that the
name Hevea brasiliensis should be kept for the Brazilian
species and Hevea Kunthiana used for the concept rep-
resented by the Venezuelan collections.
In reviewing the problem recently, Burkill (Dict.
Econ. Prod. Malay Penins. 1 (1985) 1159), stated that:
‘It would lead to much confusion were botanists at
this date to displace the... . name [ Hevea brasiliensis),
on the ground that Siphonia brasiliensis Kunth is not the
plant which everyone now calls Hevea brasiliensis; yet,
if the rules of nomenclature are followed strictly, that,
it seems, should happen. ”’
In 1986, Chevalier (in Rev. Bot. Appl. Agric. Trop.
16 (1936) 620) published a most complete review of the
historical aspects of the problem. Basing his opinions
ona study of the literature, combined with an examina-
tion of authentic collections preserved in Paris, he came
to the conclusion that the valid name for the cultivated
species of Hevea is H. brasiliensis. In his own words,
his conclusion is: ‘‘Celle [the collection] du Para [as
opposed to the Orinoco material] doit garder le nom de
Hevea brasiliensis (Willd.) Muell.-Arg. (excl. syn.
ALK.)
Although Chevalier intimated that Willdenow’s Sv
phonia brasiliensis had been validly published to denote
[ 80 |
what we today call Hevea brasiliensis, he did not enter
into a discussion of the reason why it was so according
to the Rules and why the name cannot, under any cir-
cumstances, be applied to the Orinoco material. In spite
of the fact that Chevalier’s paper constitutes the most
important contribution of the century towards a clarifica-
tion of this question, it has apparently not received the
attention it deserves. Baldwin (in Journ. Hered. 38
(1947) 54; ibid. 40 (1949) 47) accepted Chevalier’s con-
clusions, but other investigators who have recently pub-
lished on Hevea (Schultes in Bot. Mus. Leafl. Harvard
Univ. 12 (1945) 7; Seibert in Ann. Missouri Bot. Gard.
34 (1947) 305), by using ‘‘(HBK.) Muell.-Arg.’’ as
authorities for the binomial Hevea brasiliensis, have in-
dicated acceptance of the long-established belief that
Kunth’s publication of Stphonia brasiliensis was the
earliest.
Cook, in 1941, went much farther than all who had
previously discussed this problem. He _ proposed (in
Journ. Wash. Acad. Sci. 31 (1941) 46) to substitute the
new name Siphonia Ridleyana Cook for our cultivated
rubber tree. He rejected the generic epithet Hevea on
the basis of faulty reasoning and an erroneous under-
standing of the meaning of the term homonym. Even
were a new name needed, Cook’s substitute specific epi-
thet would be superfluous in view of Warburg’s Hevea
Siebert of 1900. Cook did not mention Warburg’s work
in his rather extensive discussion, nor did he indicate by
citation or by context that he was familiar with Cheva-
lier’s convincing article.
To help end the continued uncertainty in regard to
the name of such an important economic plant, and to
reiterate Chevalier’s conclusion and connect the reasons
for it with the corresponding authorizing Article of the
International Rules of Botanical Nomenclature, I shall
[ 81 ]
restate the pertinent historical facts and present as com-
plete a synonymy as possible.
In 1800, Humboldt collected material of Hevea along
the Rio Tuamini near Javita and also along the Rio
Orinoco at San Fernando de Atabapo, both localities in
the upper Orinoco basin of southern Venezuela. ‘These
collections were described by Kunth in 1825 under the
name Siphonia brasiliensis. This has been taken almost
unanimously as the first valid publication of the binomial.
In the first decade of the nineteenth century, Count
Hoffmannsegg received, presumably from the traveller-
collector F. G. Sieber, who worked in Para from 1801
to 1807, specimens of Hevea which he turned over to
Willdenow for study. There is strong reason to believe
that this material came from the lowermost course of the
Rio Amazonas. Willdenow annotated the material with
the name ‘‘Siphonia brasiliensis Willd.’’ and deposited it
in his herbarium which was preserved at Berlin-Dahlem.
These specimens represent the concept which we have
come to know as Hevea brasiliensis.
When Kunth published the name Siphonia brasiliensis,
he described the two Venezuelan collections and cited
them as the only basis for the description. He did not
cite the Brazilian material of Sieber, but he did include
in synonymy “‘Stphonia brasiliensis Willd. herb.”’ with
the following footnote: ‘‘In specimine brasiliensi a
Willdenoicum cel. Beauvois communicato (inque Museo
Lessertiano asservato) foliola multo minora, subtus pal-
lide viridia (nec albida).*” Also in synonymy, he included
‘*Siphoniae species brasiliensis Adr. de Juss. Kuphorb.
p. 40°° and ‘‘S. foliolis oblongis, acuminatis. Willd.
mss.’’ Willdenow had the habit of making such abbre-
viated descriptions on herbarium sheets or on envelopes
containing specimens, and it is entirely probable that
Kunth, who visited the Willdenow herbarium in Berlin,
[ 82 |
had copied this himself, for it seems not to have been
published elsewhere.
It is quite apparent from a thorough study of the
Kunth publication of Siphonia brasiliensis that, although
he pointed out in the footnote reproduced above that the
Brazilian material differed in several characters from the
two Venezuelan collections, Kunth himself considered
the three collections to represent the same concept. The
description, it is also apparent, was based upon the Vene-
zuelan material which, as we now know, definitely does
not represent the common cultivated plant but rather a
rare species which has, as yet, acquired no commercial
importance.
A critical examination of Kunth’s treatment discloses
an inconspicuous point which seems to have been over-
looked and which alters our interpretation of the prob-
lem. In their ‘‘Nova genera et species plantarum,”
Humboldt, Bonpland and Kunth were accustomed to
indicate names which they were publishing as their own
for the first time with a small dagger. In the preface
(loc. cit. 1 (1816) vi), they state: ‘‘Species et genera
nova signo fF indicantur.’’? Siphonia brasiliensis is not
marked with a dagger. This, coupled with their citation
in synonymy of Willdenow’s Siphonia brasiliensis (which
had been written on an herbarium sheet) would seem to
indicate that Kunth was publishing an unpublished
Willdenow name. ‘This puzzling situation is completely
clarified if, remembering the lack of the dagger in
Kunth’s publication, we refer to an article by Adr. de
Jussieu. It then becomes apparent that Jussieu validly
published Willdenow’s Stphonia brasiliensis in 1824, one
year before the appearance of Kunth’s description. In his
“De Euphorbiacearum generibus. ...’’ (1824) tab. 12,
fig. 88b, 1-6, Jussieu published a plate consisting of diag-
nostic drawings of the staminate calyx, the stamens with
[ 83 ]
EXPLANATION OF THE [ILLUSTRATION
Piare XVIII. The earliest publication of the name
Siphonia brasiliensis in Adr. de Jussieu’s “De
Euphorbiacearum generibus. .. .”’ (1824) t. 12,
fig. 38b, 1-6.
we
AD. dS. dal?
36. ALEURITES ambinusr. 4d 7. ANDA GJOMCEM,
38.A. SIPHONIA elastia. B.S. brastonets.
3
é
L 6
‘ a
Ve pore seagp'
ALVId
IIIAX
EXPLANATION OF THE ILLUSTRATION
Prare XIX. A specimen of the type collection of
Hevea brasiliensis preserved in the Paris Herbarium.
Piate SX
4
Siphoud, bone i ttehy
Wes
~ HEED. MUS. PARI
r
i/ ppherua , oe , ow
a
ete at Aweta
od Aras lous j
pv lleom,') |
255 [oe
- ce
Dede Miho snentiat, 190) -
BS
rn oe
;
the anthers and suprastaminal column, the _ pistillate
flower, the pistil and the ovary of ‘‘Siphonia brasiliensis
W. (in herbariis).’” In accordance with Article 44 of the
Rules, this constitutes valid publication, for the name
of a species is validly published if it be accompanied ‘‘by
a plate or figure with analyses showing essential charac-
ters; but this applies only to plates or figures published
before January 1, 1908.”’
When Mueller transferred the specific epithet from
Siphonia to Hevea in 1865, he was probably unaware of
the discrepancy between the Orinoco and the Amazon
collections—and this in spite of Baillon’s insistence on
that point in 1858. In making the new combination,
Mueller based it on ‘‘Stphonia brasiliensis Kunth in
Humb. et Bonpl. Nov. Gen. et Spec. 7, p. 171°” and
cited “*S. Kunthiana Baill.’ as a synonym. It is signi-
ficant, however, that the Brazilian material (‘‘In Brasilia
paraénsi, Hoffmansegg in hb. Willd. fol. 17986. p. 1°’)
was cited before the Venezuelan collections. It is because
the author of this combination, apparently unaware of
Jussieu’s paper, attributed the earliest publication to
Kunth that the author citation of Hevea brasiliensis has
been erroneously written as ‘‘(HBK.) Mueller-Argovi-
ensis’’ by almost all taxonomists.
In summary, we may say that the proper and valid
name of the cultivated rubber tree is
Hevea brasiliensis (Willd. ex Adr. de Juss.)
Mueller-Argoviensis in Linnaea 84 (1865) 204.
Its synonymy is as follows:
Siphonia brasiliensis Willdenow ex Adr. de Jussieu
Euphorb. Gen. (1824) t. 12, pl. 88b., fig. 1-6. Mon
Siphonia brasiliensis HBK. Nov. Gen. et Sp. 7 (1825)
wee
Hevea janeirensis Mueller-Argoviensis in Martius F].
Bras. 11, pt. 2 (1874) 706.
Hevea Sieberi Warburg Kautschukpf. (1900) 82, fig.
[ 85 |]
Hevea Randiana Huber in Bol. Mus. Goeldi 4 (1906)
656.
Hevea brasihensis var. stylosa Huber (loe. cit. 4 (1906)
640.
Hevea brasihensis var. janeirensis (Muell.-Arg.) Pax
in Pflanzenr. 4 (1910) 121.
Hevea brasiliensis var. Randiana (Huber) Pax loe. cit.
4 (1910) 123.
Hevea brasiliensis mut. Granthami Bartlett in Bot.
Gaz. 84 (1927) 200.
Hevea Granthami Bartlett loc. cit. 84 (1927) 200,
nomen alt. anteriori.
Hevea brasiliensis forma typica Ducke in Arch. Jard.
Bot. Rio Janeiro 6 (1988) 55.
Hevea brasiliensis var. subconcolor Ducke loc. cit. 6
(1988) 55.
Hevea brasiliensis torma subconcolor Ducke in Arch.
Inst. Biol. Veg. Rio Janeiro 2 (1985) 224.
Hevea brasiliensis forma Randiana (Huber) Ducke
loc. cit. 2 (1985) 224.
Siphonia Ridleyana Cook in Journ. Wash. Acad. Sci.
31 (1941) 46.
Siphonia janeirensis (Muell.-Arg.) Cook loc. cit. 31
(1941) 61.
The varietal names which follow, included by Seibert
(loc. cit. 805-806) as synonyms of Hevea brasiliensis, are
here omitted pending additional studies which may pos-
sibly indicate that they are distinct concepts.
Hevea brasiliensis var. angustifolia Ule in Tropen-
pflanz. Beiheft. 6 (1905) 8.
Hevea brasiliensis var. latifolia Ule loc. cit. 6 (1905) 8.
Hevea brasiliensis var. acreana Ule in Engler Bot.
Jahrb. 50 (1914) 14.
For his friendly counsel in this as in many other prob-
lems which have arisen in my botanical research, I am
deeply grateful to the late Mr. Charles A. Weatherby
of the Gray Herbarium of Harvard University.
[ 86 ]
THE IDENTITY OF UCUQUI
BY
Joao Murea Pires' and Ricuarp Evans ScHULTES?’
ONE of the results of recent field work in the upper
Rio Negro basin of Brazil has been the identification of
a useful plant of that area—the ueuqui. The fruit of this
tree has an edible and delicious mesocarp and is an impor-
tant part of the diet of the native peoples of the region.
Investigation has shown that the uweuqui is an unde-
scribed species of the sapotaceous genus Pouteria. It is
altogether fitting that, in publishing a description of this
food plant, we employ as a specific epithet the common
name which refers exclusively to this species over the
greater part of its range.
Pouteria Ucuqui is immediately set apart from all other
species of the genus by the excessively developed disk
which surrounds the ovary.
Pouteria Ucuqui Pires & Schultes sp. nov.
Arbor enormis, usque ad centum viginti pedes alta,
radicibus tabularibus, trunco columnari usque ad _ tres
pedes in diametro, cortice crasso, molli, extus atrobadio
‘Chief, Section of Biology, Instituto Agronémico do Norte, Belém
do Para, Brazil.
Botanist, Bureau of Plant Industry, Soils, and Agricultural Engi-
neering, Agricultural Research Administration, U.S. Department of
Agriculture; Research Fellow, Botanical Museum, Harvard Univer-
sity; Collaborator, Instituto Agronémico do Norte.
[ 87 ]
et intus sanguineo cum latice albo aquosoque. Ramuli
juniores, inflorescentiae, petioli et foliorum nervi indu-
mento ferrugineo-pulverulento vel ferrugineo-furfuraceo
obtecti. Folia alterna, bene coriacea, elliptica, basi et
apice acuta vel obtusiuscula, plerumque cum acumine
7-10 mm. longo, margine integra, 11-20 ecm. longa,
5.5-10 em. lata, supra nitidula, subtus opaca (statu bene
juvenili ferruginea), minutissime ferrugineo-pilosula;
costa leviter striata, supra prominens, subtus valde elevata
et robusta; nervi secundarii utrinque octo ad quindecim,
supra leviter insculpti, subtus valdissime prominentes ;
venulae superficie utraque paulo impressae. Petiolus
elongatus, siccitate striatus et leviter rugosus, saepissime
3.5-5.5 em. longus, 2-83 mm. in diametro. Inflorescen-
tiae axillares, valdissime congestae, fasciculatae. Flores
mordaciter fragrantes, flavo-virides, gemmationis statu
longe persistentes sed flores ipsi fugaces. Sepala quin-
que, imbricata, subrotundata, plerumque 2 mm. longa,
utrinque minutissime adpresso-pilosa, intus demum gla-
brescentia. Petala quinque, per + vel $ longitudinis par-
tem connata, imbricata, oblonga, apice acutiuscula vel
obtusa, margine integra, plerumque 3 mm. longa, 1 mm.
lata, extus pilosiuscula sed demum glabrescentia, intus
subglabra, tubo vulgo 0.5-1 mm. longo. Stamina quin-
que, petalis opposita, filamentis vix 1 mm. longis; an-
therae apice acutae, basi rotundatae, connectivo mediano
crasso, loculis duobus, sublinearibus, rimis oppositis, de-
hiscentibus. Staminodia petalis alterna, apice acuta, 1
mm. longa vel minora, aliquando inconspicua. Ovarium
biloculare, minutissimum, pilosum, disco valde hispido,
circiter 0.5 mm. alto circumdatum; stylus circiter 1.2
mm. longus, pilosus vel glabrescens, cum stigmatibus
inconspicuis. Fructus maximus, 9-13 cm. longus, 5-7
cm. in diametro, monospermus; semen magnum, pler-
umque 7-10 cm. longum, 3.5 cm. latum, 3 em. in di-
[ 88 |
ametro, cum testa crustacea, nitida, plusminusve 2 mm.
crassa, area umbilicali magna.
Arbor a regionis typicae brasiliensis incolis weuqué (in
lingua nheenagtti), puch-pee-d (in lingua tukanorum tri-
bus); a regionis colombianae incolis 06-le-da (in lingua
kuripakanorum tribus), /a-he-pa (in lingua mirahorum
tribus) appellatur.
CoLLECTIONS EXAMINED:
Brazit: Estado do Amazonas, Upper Rio Negro, Igarapé Uaba,
opposite mouth of Rio Xié. ‘‘Immense tree, 110 feet tall, with wide-
spreading crown. Large buttress roots up to 6 feet. Tree columnar,
24 to 3 feet in diameter. Bark dark brown, shaggy externally, red
internally, soft, }—? inch thick. Latex sparse, white. Wood of medium
hardness, compact, white. Flowers in extremely dense clusters, fall-
ing easily. Sepals greenish; petals white; anthers brown, apparently
odourless. Common name: ucuqut.’’ January 5, 1948, Richard Evans
Schultes & Francisco Lopez 9553 (Tyre in Herb. Gray; Dupticatr ryprs
in Herb. Inst. Agron. Norte (Belém do Parad); U.S. Nat. Herb. ;
Kew; Econ. Herb. Oakes Ames).—Rio Issana basin, between Rio
Aiari and Rio Caiari (Uaupés), near Serra Tunui. “‘Arvore 25 m.
40 em. Frutos estimados como alimento indigena. Ucuqut.’? October
13, 1945, Ricardo de Lemos Frées 21388.—Rio Negro, near Uaupés
(Sao Gabriel), Serra de Uanari ‘‘Ucuqui. Arvore muito grande. Latex
branco, pegajoso, pagina inferior da folha revestida por leve indumento
escamoso ferrugineo, fruto amarelado.’’ October 26, 1947, Joao Murca
Pires 773.—Same locality. ** Ucuqué. Arvore muito grande. Latex
branco, pegajoso, pagina inferior da folha revestida por leve indumento
escamoso ferrugineo; fruto amarelo.’’? October 30, 1947, Joao Mur¢a
Pires 799.—Same locality. *‘ Ucuqut. Arvore muito grande no alto da
serra. Floras quasi brancas (botoes).’? November 17, 1947, Joao Mur¢a
Pires 1165.—Rio Negro, near confluence with Rio Uaupés. ‘‘ Ucuqut.
Arvore muito grande. Madeira dura. Latex branco, pegajoso. Pagina
inferior da folha revestida por leve indumento escamoso, ferrugineo,
amarelado.’’ November 5, 1947, Joao Murga Pires 833,— Rio Uaupés,
Serra Uapici. ‘‘Ueuqué. Arvore grande.’’ November 17, 1947, Joao
Murca Pires 1150.—Rio Negro, near Uaupés (Sao Gabriel). ** Ucuqui.
Arvore platinhas novas, colhidas sob a arvore.’’ October 30, 1947, Joao
Murga Pires 792.—Rio Uaupés, Ipanoré **Puh-pid. Arvore grande.’’
November 15, 1947, Joao Murca Pires 1057.—Rio Uaupés, Taracua.
[ 89 ]
**Ucuqui, (lingua geral), Puh-pid (Tukano), November 8, 1947, Joao
Murca Pires 905.—Rio Uaupés, Serra Uapici. ** Ucuqui. Arvore
grande.’’ November 17, 1947, Joao Murca Pires 1149.—Rio Negro,
Sao Gabriel. “‘Coqui. Enormous tree. Buds brownish. Fruit edible.”’
November 20-25 1947, Richard Evans Schultes & Francisco Lépez 9184.
—Rio Negro, Sao Gabriel. “‘Ucuquéi. Enormous tree.’’ January 14,
1948, Richard Evans Schultes & Francisco Lopez 9618.—Middle Rio
Negro basin, Rio Curicuriari. “‘Ucuqui.’’? Richard Evans Schultes &
Francisco Lopez 9710.—Rio Negro, Ipanoré, Caatinga forest beyond
town. ‘‘Coqut. Enormous tree. Fruit edible.”” November14—15, 1947,
Richard Evans Schultes & Joao Murca Pires 9096.
Cotomsta : Comisaria del Vaupés, Rio Negro, opposite Piedra del Co-
cuy. ‘Enormous buttressed tree, 110 feet tall, diameter 3 feet. Flowers
greenish yellow. Bark shaggy, brown. Fruit edible.’’ December 28,
1947, Richard Evans Schultes & Francisco Lépez 9484.—Rio Guainia,
near Sejal. ““Tree 100 feet tall, buttressed. Kuripaka name: 06-/e-da.”’
June 1948, Richard Evans Schultes & Francisco Lépez 10058a.—Com-
isaria del Amazonas, Rio Caqueté, near La Pedrera. ‘* Enormous
tree. Fruit edible. Mirafia name: kd-he-pa.’’ July 15, 1948, Richard
Evans Schultes & Francisco Lopez 10215,
VenezurLa: Territorio del Amazonas, Rio Negro, base of Piedra
del Cocuy. ““Enormous tree 120 feet tall, 24 feet in diameter, slightly
buttressed. Bark brown, shaggy, soft, scarlet within. Latex sparse,
white. Buds greenish yellow.’’ December 24, 1947, Richard Evans
Schultes & Francisco Lopez 9458.
Pouteria Ucuqui is very common in the northwestern-
most part of the Brazilian State of Amazonas along the
upper Rio Negro, from 'Tapurucuara (Santa Isabel) up-
stream, and along its affuents: the Uaupés, Issana,
Tikié, Curicuriari, Dimiti, and probably many others.
It is known to occur in Colombia in the Rios Guainia
(constituting the source of the Rio Negro) and on the
Rio Caqueta. It has also been found on the Japura in
Brazilian territory and has been reported from the Rio
Solimoes (Le Cointe, P. ‘‘Arvores e plantas uteis’’ (1934)
457). According to reliable reports, it occurs in certain
places far into Venezuelan territory along the lower
course of the Rio Guainia and on the Casiquiare itself.
[ 90 |
In this entire area, the plant—an enormous, heavily
crowned tree which fruits profusely once a year—is found
in abundance in the virgin forest on high, well drained
soil. [t is also often encountered in a state of apparent
cultivation, since, in clearing for house sites, the tree has
been spared. When completely ripe, the thick, fleshy
mesocarp of the fruit, which resembles that of the avo-
cado (Persea americana Mill.) to a striking degree, is
very palatable. When green, however, it is full of latex
and is extremely sticky.
Pouteria Ucuqui is known in the entire Brazilian part
of its range by the name weugu?, a word originating from
the Lingua Geral or Nheengatti language which is spoken
widely in the State of Amazonas, and especially in the
Rio Negro. The Tukano Indians of the Rio Uaupés re-
fer to the tree as puch-pee-d (the ch being soft as in the
German ?ch). In Colombia, the native name for Pouwteria
Ucuqui amongst the Kuripaka Indians of the Rio Guainia
is 00-le-da; and amongst the Miranias of the Rio Caqueta
(La Pedrera), hd-he-pa. The tree is called yuew by the
Spanish-speaking population of the Venezuelan town of
San Carlos on the uppermost Rio Negro.
The term weuqu?, so far as we have been able to ascer-
tain, refers exclusively to Pouterta Ucuqui and should
not be confused with weuquirana (i.e., ‘false weuqut’’ ),
a name widely applied in the same area to the sapotaceous
Ecclinusa sanguinolenta Pierre (EH. Balata Ducke)—also
‘alled abiwrana—a tree which is actively exploited as the
source of a type of balata. The two trees are completely
distinct from all points of view.
Pouteria Ucuqut is a very tall and robust tree, usually
with comparatively large buttresses at the base; the low-
est branches are at a great height from the ground. These
conditions make the study and collection of herbarium
material rather difficult, especially since the flowers,
[ 91 ]
which are borne on the branchlets, are very minute and
‘annot be seen from the ground, even with binoculars.
Dr. Adolpho Ducke, who has visited the upper Rio
Negro basin several times, was unable to collect flowering
material of weuqui, although he devoted special attention
to this tree which he believed represented an undescribed
species. Without flowering material, however, there was
some question as to even its generic affinities.
When we began our work in the upper Rio Negro in
late 1947, Dr. Ducke counseled us to try to find flower-
ing material which would settle definitively the identity
of ueugui. With this in mind, we studied a number of
individuals over a wide area. It was almost always neces-
sary to use the balatero’s climbing-irons (employed dur-
ing the extraction of balata from species of Manilhara),
since the girth of the tree usually prevented the use of
the ‘‘peconha”’ (a band of pounded bark which the In-
dians place on the feet to aid them in climbing). The
expertness of our assistant, the late Francisco L6pez, in
the use of the climbing-irons greatly increased the num-
ber of trees which we could study.
During October and November, this work was carried
on principally by Murea Pires. At first, sterile specimens
only were obtainable. Then, gradually, we began to se-
cure material which was in bud. No flowers, however,
were found, and we later learned that the species develops
the flower with extreme slowness, persisting in bud some-
times for three or four months. Many experiments were
tried to force the buds to open: sun, artificial heat, soak-
ing in hot and warm water; but all these efforts failed.
It is interesting in this connexion to note how little
the natives know about the life-history of this species,
even though it is one of their common and useful plants.
Repeated questioning on our part brought forth the most
divergent and amusing remarks concerning the flowering
[ 92 ]
of weuqut. Many, when asked during what month the
tree flowers, replied that they had never seen it flower
and that, therefore, it did not flower. Others insisted
that the tree blooms during the night and immediately
drops all the flowers. This second ‘‘explanation”’ is, in
a way, ingenious, because, as we later witnessed, the forest
floor under an weuqui tree, where there had even the day
previously been no indication of blossoms, would sud-
denly be covered with literally hundreds of thousands of
flowers or parts of flowers. Our studies showed that, in
spite of the very long period of aestivation, the flowers
are almost ephemerous in Pouteria Ucuqut.
During the last few months of 1947, Murea Pires
revisited trees in bud several times in an attempt to col-
lect fertile specimens. When he returned to Belém,
Schultes and Lopez continued the search. In late De-
cember, it was noted that the buds on a number of trees
previously examined were swelling rapidly, in spite of
the fact that for at least a month previous there had been
no appreciable alteration in their size.
On January 5, we found our first flowering tree. It
was very early in the day, about six o’clock in the morn-
ing, when our attention was called to the tree by the
patter of falling flowers. Upon climbing the tree, we
noted a very strong aromatic and pungent odor. It is
significant to note that the buds never had a fragrance.
There were innumerable large bees and several other
kinds of smaller insects visiting the flowers even at this
early hour. Later, we encountered other trees in flower,
but the pronounced odor was not noted. It is possible
that the strong, aromatic fragrance is given off only for
a short period during the very limited blossoming time.
As accurately as we could calculate, open flowers per-
sist for no longer than three days on an individual tree
and usually are much shorter lived.
[ 93 |
EXPLANATION OF THE ILLUSTRATION
PLrate XX. Pourerta Ucugut Pires & Schultes. A,
flowering branch, about one half natural size. B,
fruit. C, seed. D, flower. E, dissection and view
of interior of corolla. F, dissection of flower show-
ing disk around ovary. G, dissection of flower
showing pilosity around ovary. H, anther. Each
drawing has a millimeter scale to indicate magni-
fication.
Drawn by Perera Fitno
Qf Instituto Agronomico do Norte
PRATE XX
POUTERIA
EXPLANATION OF THE ILLUSTRATION
Prare XXII.) Pourerta Ucugut Pires & Schultes.
Basal portion of trunk of type tree, showing the
buttress roots.
Photograph by R. EB. Scuvurres
4 |
’
‘
PLATE
<a
om
ee
=
ede ad
EXPLANATION OF THE TLLUSTRATION
Prare XXIT. Pourerta Ucugut Pires & Schultes.
Inflorescences from branchlets of type tree.
Photograph by R. BK. Scuuvres
PLATE XNII
KXPLANATION OF THE ILLUSTRATION
Prare XXIII. Pourerta Ucugur Pires & Schultes.
Flowers from type tree.
Photograph by R. EK... Scuvurres
PLATE XXITI
A century set (i.e. one hundred herbarium specimens)
was collected from the type tree for distribution by the
Gray Herbarium of Harvard University. Floral material
was also obtained from the same tree and preserved in al-
cohol and glycerine for use in drawing up the description.
An interesting Indian use of weuquit, discovered by
Froées along the Rio Japura, is the manufacture of toy
whistles from the seeds. All of the interior of the seed
(the cotyledons) is removed and a number of holes, over
which the fingers are placed, are made. In popular medi-
cine in Amazonian Brazil, the fruit is sometimes em-
ployed as a vermifuge (Le Cointe loc. cit.).
The earliest and, in fact, the only reference of an ex-
tended nature to ucuqui which has come to our attention
is Wallace’s interesting discussion of the plant (Wallace,
A. R. ‘‘A narrative of travels on the Amazon and Rio
Negro”’ ed. G. T. Bettany (1889) 243-244). Because of
its historical significance, we quote it in full. ‘‘Of a
passing Indian {on the Rio Uaupés] I bought a basket
of Ocoki, and some fish. The Ocoki is a large pear-
shaped fruit, with a hard thick outer skin of almost
woody texture, then a small quantity of very sweet pulpy
matter, and within a large black oval stone. The pulp
is very luscious, but is so acrid as to make the mouth
and throat sore, if more than two or three are eaten.
When, however, the juice is boiled it loses this property ;
and when made into mingau with tapioca, is exceedingly
palatable and very highly esteemed in the Upper Rio
Negro, where it is abundant. It takes at least a peck of
fruit to give one small panella of mingau.
“On the next day, the 10th in the afternoon, the In-
dians all suddenly sprang like otters into the water, swam
to the shore, and disappeared in the forest. ‘Ocoki’ was
the answer to my inquiries as to the cause of their sud-
den disappearance ; and I soon found they had discovered
[ 95 |
an ocoki-tree, and were loading themselves with the fruit
to satisfy the cravings of hunger, for the Indian’s throat
and mouth seem invulnerable to all those scarifying sub-
stances which act upon civilised man. The tree is one of
the loftiest in the forest, but the fruit falls as soon as
ripe, and its hard woody coating preserves it from injury.
Baskets, shirts, trousers, ete,, were soon filled with the
fruit and emptied into the canoe; and I made each of
the Indians bring a small basketful for me; so that we
had ‘mingau de ocoki’ for three succeeding mornings. ’’
[ 96 ]
THE CORRECT NAME OF THE YAUPON
BY
RicHarp Evans ScCHULTEsS’
DvriING a recent conversation with me regarding the
nomenclature of the yaupon, Professor Oakes Ames
‘alled my attention to a precious remark which Dillenius
once wrote in a letter to Linnaeus (Smith, J. E.: ‘SA
selection of the correspondence of Linnaeus and other
naturalists’’ 2 (1821) 96). *‘We all know the nomencla-
ture of Botany to be an Augean stable, which C. Hoff-
man, and even Gesner, were unable to cleanse.”” Noth-
ing could be more expressive of the confusion which one
encounters in the synonymy of this holly than the term
** Augean stable. ””
There has long been a need for a thorough discussion
of the correct name of the yaupon, the source of the
black-drink once so commonly used as a ceremonial emet-
ic and stimulant amongst Indians of the southeastern
part of the United States.
The monographer of the Aguifoliaceae, Loesener
(Monogr. Aquifol. in Nova Acta Acad. C. L. C. G.
Nat. Cur. 78 (1901) ) employed the binomial J/ex caro-
liniana (Iuam.) Loes. Most modern taxonomic treat-
ments, however, use Ilex vomitoria [Soland. in] Ait.
' Botanist, Bureau of Plant Industry, Soils, and Agricultural Engi-
neering, Agricultural Research Administration, United States Depart-
ment of Agriculture; Research Fellow, Botanical Museum, Harvard
University. This contribution is part of a study of New World narcotic
and stimulant plants carried out as a Guggenheim Fellow in Botany.
[ 97 ]
Notwithstanding this tendency in botanical works, a
most bafHing confusion and uncertainty. still exists
throughout the numerous chemical, pharmacological,
anthropological and ethnobotanical publications and
hampers understanding and accuracy. In view of the
lack of any adequate review of the taxonomic and no-
menclatorial history of this plant since 17538, it has seemed
advisable to present the following notes in an attempt to
set forth the reason why the correct, though not the
earliest, name is [/ex vomitoria and to discuss the origin,
status and significance of the synonymy of this binomial.
Ilex vomitoria [Solander in] Aiton Hort. Kew.
Led. 1] 1 (1789) 170.
I. Cassine B Linnaeus Sp. Pl. 1 (1758) 125.
Cassine Paragua Miller Gard. Dict. [ed. 8] (1768)
Cassine no. 2. Non Cassine Peragua Linnaeus Sp.
Pl. Led. 2] (1762) 884. Non Cassine Peragua Linnaeus
Mant. 2 (1771) 220.
C. caroliniana Lamarck Dict. 1 (1783) 652, pro parte,
I. Cassine Walter F 1. Carol. (1788) 241.
I. hgustrina Jacquin Collect. 4(1790) 105; Icon. Rar.
2 (1793) 9, t. 310.
Casine yapon Bartram Travels (1791) 259, nom. nud.
I. floridana Lamarck Ill. 1 (1791) 856.
I. Cassena Michaux FI. 2 (1808) 229.
I. religiosa Barton Fl. Virg. (1812) 66.
? Cassine ramulosa Rafinesque in Robin FI. Lud.
(1817) 110.
Mierophyllus Cassine (Walt.) Rafinesque Med. Bot.
2 (1830) 8.
? Cassine amulosa Rafinesque Med. FI. 2 (1880) 205,
nom. sphalm.
Ageria Cassena (Michx.) Rafinesque Sylv. Tellur.
(1838) 47.
[ 98 |
? Emetila ramulosa Rafinesque in S. Watson Bibl.
Ind. N. Am. Bot. (1878) 158, nom. in syn.
Cassine yaupon Gatschet in Trans. St. Louis Acad.
Sci. 5 (1888) 88, nom. nud.
I. Peragua (.) Trelease in Trans. Acad. Sci. St.
Louis 5 (1889) 346, in obs.
I. caroliniana (Luam.) Loesener in Bot. Centralbl. 47
(1891) 163.
I. vomitoria [Soland. in] Aiton var. Yawhkeyii Tarbox
Some Nat. Hollies in Brookgreen Gardens (1944) 19,
sine diagn. lat.
Cassine vonutoria Swanton in Bull. Bur. Am. Ethnol.
137 (1946) 284, nom. nud.
Perhaps the earliest widespread reference to the yau-
pon in a botanical publication was made by Bauhin and
Cherler in their encyclopaedic ‘‘Historia plantarum uni-
versalis”’ 8 (1651) 631. No description of the plant was
given. The title, ‘‘Herba Cassiana, famen sitimque re-
tardans,’’ of the brief discussion, which reported the use
of the infusion of the leaves as a stimulant, referred to
the hunger-allaying properties of the shrub. This refer-
ence is an elaboration of an earlier report by Bauhin
(Pinax (1628) 170), based upon what appears to be the
first account of the black-drink (Nufiez Cabeca de Vaca
‘*Relaci6n y comentarios. ...*’ (1542) cap. 26), herein
reproduced (Plate X XIV).
Later, Leonard Plukenet described the species as Cas-
sine vera Floridanorum Arbuscula baccifera Alternati
Ferme facie, foliis alternatim sitis, tetrapyrene (‘*‘Opera
omnia botanica’’ 8 (1691) 40) and published what is ap-
parently the earliest illustration (loc. cit. 4 (1692) t. 876,
f. 2). No mention seems to have been made of the use of
the plant.
It is to Mark Catesby (‘“The natural history of Caro-
lina, Florida and the Bahama Islands’’ 2 (1754) 57) that
we are indebted for the first definite and convincing iden-
[ 99 ]
tification of the source of the black-drink which, for 285
years previously, had been attracting the attention of
travellers. his was accomplished through Catesby’s
charming plate and his description of Plukenet’s Cassine
vera Floridanorum Arbuscula baccifera Alternati ferme
Jfacie, folis alternatim sitis, tetrapyrene, together with a
very lengthy account of the properties and ceremonial
uses of the plant.
The earliest tenable post-Linnaean name for the yau-
pon is found in volume one of the first edition of Aiton’s
‘*Hortus Kewensis’’ (1789). It was here named Ilex
vomitoria. The specific epithet would indicate that Solan-
der, who described the plant, was familiar with its renown
as an emetic, although he made no mention of this in
his notes. The original description is short: ‘‘I. foliis
alternis distantibus oblongis obtusiusculis crenato-serra-
tis: serraturis muticis.”’
It will be noted that Miller’s Cassine Paragua, (1768)
referring, without any doubt, to the concept now under
consideration, antedates [lea vomitoria. We must inves-
tigate the reasons for the untenability of Miller’s bino-
mial, because of the almost unbelievable confusion of
his specific epithet and Linnaeus’ several concepts—all
named Cassine Peragua or the orthographic variant C.
Paragua. Notwithstanding the unavailability of the epi-
thet Peragua, the combination under J/ex has been
made. We must realize, however, that the author did not
urge its acceptance. Trelease, although he accepted [lew
Cassine Walt. as the correct name for the yaupon holly,
wrote in a footnote (in Trans. Acad. Sci. St. Louis 5
(1889) 346, im obs.): ‘‘ An effort to improve on the no-
menclature of this species, unless it is called J. Cassine
BL. Sp. (1758) 125, would probably cause it to stand as
I. Peragua (.)=Cassine Peragua L. Mantiss. (1771),
ii, 200.”
[ 100 |
In 1762, Linnaeus (Sp. Pl. [ed. 2] 1 (1762) 384) pub-
lished Cassine Peragua tor an Old World concept.
He cited several sources which indicate that the concept
was unquestionably an Old World one. Although one
of these sources (Hortus Cliffortianus (1737) 72) attrib-
uted the plant both to the Cape of Good Hope and to
Carolina, it is clear that it was not being confused with
the concept now known as JIlea vomitoria which was
discussed separately (loc. cit. 40) and was attributed to
“Carolina A mericae.””
Cassine Peragua was published again by Linnaeus
nine years later (Mant. 2 (1771) 220). In this second
Linnaean publication, it referred not to the Old World
concept of the earlier (1762) Cassine Peragua but to
several different concepts.
Linnaeus’ Cassine Peragua of 1771 has hitherto been
considered by some taxonomists as a synonym of Ilex
vomitoria, either wholly or in part. There has been much
uncertainty as to the exact meaning of this name, be-
‘ause, In the Linnaean Herbarium, there are two distinct
American plants under the binomial. Loesener (in Engler
Bot. Jahrb. 28, 2 (1900) 154-155) considered Cassine
Peragua L. of 1771 as a nomen nudum. Fernald and
Schubert (in Rhodora 50 (1948) 169) hold it to be a nomen
amhiguum and a nomen confusum and suggest that it be
permanently rejected. Photographs of the two specimens
indicate that neither is referable to Ilex vomitoria; both
are species of Viburnum (Fernald and Schubert loc. cit. ).
Therefore, whether Cassine Peragua L. of 1771 be re-
jected or not, there is no need of considering further the
name in any discussion of the synonymy of lea vomi-
toria. 'Trelease’s combination, I/evw Peragua is untenable
since it is based on a homonym.
Cassine Paragua Miller (Gard. Dict. [ed. 8] (1768) Cas-
sine No. 2), on the contrary, is very definitely referable to
[ 101 ]
EXPLANATION OF THE ILLUSTRATION
Prare XXIV. The earliest European account of
the use of the black-drink was written by Nufez
Cabeca de Vaca and published in his ‘‘Relacién y
** of which there have been
commentarios....
several editions. The illustration is a photograph
of the account as it appears in the first edition of
1542. Of this rare and fascinating book only two
copies are known: an imperfect copy preserved in
the British Museum; and a perfect one in the New
York Public Library, through the kind coopera-
tion of which institution the photograph herein
reproduced was made available.
PLATE XXIV
thrrafe embosrachan covn bumoy 4 quato tiencn
pozrcl, beven tambien otra cola q facan oelas bojas
oclosarboles como de enjzina y tucttanla en puos bo
tesal fuego-y oefpucs Gis tienen toftadabinchen el
bote dc agua tafliloticnen fobze el fuego~% quando
baberuido dos vezes echante en yna vafiat cfta cn
friandola con media calabaca:y quando efta con mus
cha efpuma beuen la tan caliente quanto pueden {uf
frirsy oefdequelafacan vel bote hafta que la beuen e+
{tan dando boxes oi3tendo que quien quierebeuer.¥
quandolas mugeres oxen eitas boxes luegofe paran
fin ofar fe mudar-y apng eften mucbocargadas no o-
fan bajer otra cofa.yfiacafo alguna vellas fe mucue
lavefbonrran tla oan oe palos-ycon muy gran end
jo derraman clagua quetienen para beucrszla quean
beuidola toznan alancarloqualellos bazeu muy lige
ramente tfin pena algiéa.fa razon defta coftumbze 02
ellos-y vizen.due fi quado ellos quieren beuer aque
llaagualas mugeresfe mueuen de donde les toma la
bos queenaquella agua feles mete enel cuerpo vna
cofumalay G dédea pocoles base mozr.y todo el tie
po gelagua eftacoziendoa de eftar elbote atapado
¥ fia cafo efta vefatapadoyalguna muger pafialove
rrama y nobeuen masoe adilaagua-esamarils.y e
ftan beuiendola tres viasfin comer: y cada dia beue
cada ynoarroba y media della. E qndolas mugeres
eftacofu coftumnbze no bufcan ve comer mas Oe para
fifolas pozg ninguna otra perfonacomedelo que ella
trac.Enel tiépo dq afi eftaug entre cltos vi vna diablu
EXPLANATION OF THE ILLUSTRATION
Prater XXV. The plant which was the source of
the black-drink was definitely identified in 1754
by Mark Catesby, as evidenced by this beautiful
illustration from his ““The natural history of Caro-
lina, Florida and the Bahama Islands’? 2 (1754)
57.
PLATE: X XV
Cassena de
EXPLANATION OF THE ILLUSTRATION
Pirate XXVI. The discussion of the black-drink
accompanying Catesby’s plate.
PEATECN SY |
Caffena vera Floridanorum, Arbufcula baccifera Alaterni facie, foliis alternatim
fitis, tetrapyrene.
T ‘HIS Shrub ufually rifes from the grotind with feveral
ftems, to the beige of twelve feet, {hooting into man
upright, flender, {tiff branches, covered with whitith {moo'
bark, fet alternately with fmall evergreen ferrated leaves, re-
fembling thofe of A/aternus. Its flowers are fmall, white, and
row promifcuoufly amongft the leaves, and are fucceeded by
mall fpherical berries on fhort footftalks: thefe berries turn
red in Oéfober, and remain fo all the winter, which, with the
green leaves and white bark, make an elegant appearance. But
the great efteem and ufe the American Indians have for it,
gives it a greater character. They fay, that, from the earlieft
times, the virtues of this Shrub has been known, and in ufe
among them, sik in the manner they now do it, which is
after having dried, or rather parched the leaves in a porrage-
pot over a flow fire, they keep it for ufe: of this they prepare
their beloved liquor, making a ftrong decoction of it, which
they drink in large quantities, as well for their health as with
great guft and pleafure, without any fugar or other mixture ;
yet they drink and difgorge it with eafe, repeating it very often,
and {wallowing many quarts. ‘They have an annual cuftom in
the {pring of drinking it with ceremony ; the town having no-
tice from the King or principal, the inhabitants affemble at the
Town-houfe, having previoufly by fire purged their houfes of
all their old furniture, and fupplied them with new, the King
is firft ferved with a bowl or conch-fhell, never ufed before, of
this emetick broth, by the next to him in eminence, and he by
the next is ferved ; and fo on, till he comes to the women and
children. They fay it reftores loft appetite, {trengthens the
ftomach, giving them agility and courage in war, &e, It
grows chiefly in the maritime parts of the country ; from
whence thofe Indians {upply the Mountain-Indians with it ; car-
rying on the like trade with it in Florida, as the Spaniards do
with the South-fea tea from Paraguay to Buenos-ayres ; Florida
being in the fame latitude north, as Paraguay is fouth: and
obferving, by comparing the leaves of both, no apparent dif-
ference in them, induces me to believe they are both the fame
Plant. In South-Carolina it is called Cafena. In Virginia and
North-Carolina it is known by the name of Yapon; in the latter
of which places it is as much in ufe te the white people
as among the Indians, at leaft among thofe who inhabit the {ea-
coafts.
V-0O'L. IE,
Pluk. Mant.
Cer Arbriffeau séleve ordinairement a2 la hauteur de douze
pits, avec plufieurs fgets gui fe divifent en plufieurs branches
menues, roides, couvertes d'une écorce liffe & blanchdtre, & garnics
de petites feuilles dentelées, toujours vertes, rangées alternative-
ment, & femblables 2 celles de I'alaterne. Ses fleurs font petites,
blanches, & viennent ca & la parmi les feuilles. Al leur fucvede de
petites bayes fpbériques, atiachées a des pédicules courts: ces layes
deviennent rouges au mois d’'Octobre, & demcurent en cet état tout
I’'Hiver, ce qui, joint au verd des feuilles & 2 la blancheur de I'é-
corce, produit un effet tres agréable; mais le mérite de cet Arbriffcau
eft encore plus relevé par I'e/time que les Américains ont pour lui, &
Vufage qu'ils en font : ils difent que fes vertus ont été connues parmi
eux depuis les prémiers temps, é quils sen font fervis en le prépa-
rant de la méme maniere qu'ils le font aujourd Lui: ce qui fe fait en
Jechant, ou plutét en piles fur un petit feu dans un pot les feuilles
que l'on veut garder: ils font de ces feuilles une forte d:cocticn,
gui eft leur breuvage favori, dont ils boivent une grande quan-
tité autant par plaifir & par goiit, que pour leur fanté: tls n'y
milent ni fucre ni autre chofe; & ils en avalent plufeurs pintes,
gu'ils vomiffent avec la méme facilité, ce qu'tls repetent fouvent.
Ils ont une coiitume annuelle au Printemps de boire cette liqueur en
cérémonie: la ville étant avertie par Je Roi ou le Chef, les babitans
Saffemblent 21a maifon de ville, apres avoir purgé leurs matfons
de tous leurs vieux meubles, €& les avoir garnies de nouveaux uten-
Siles: le Roi regoit le prémier dans un vutffeau cu une coguille, qui
n'a jamais fervi, cette liqueur émétique des mains du plus qualifié
apres lui: celui la of fervi par un autre; & on va ainfi de futte
Jufguaux femmes & aux enfans. Ils difent que ce bouillon retablit
Vappetit, fortifie l'eflomac, & leur donne de l'agilité & du courage
@ la guerre, &c. Il croit le plus communément dans les cndrotts
du pays qui fut vers la Mer; & les Indiens, qui les habitent, en
Sourniffent 2 ceux des Montagnes : ils en font le méme commerce a
Ja Floride, que les Efpagnols font du thé de la Mer du Sud, depuis le
Paraguai ju/gu'd Buenos-Ayres. La Floride éfant dans la méme
latitude au Nord, que Je Paraguai eff au Sud, & ne trowvant aucune
différence entre les feuilles de fun & de l'autre, je fuis porté a
croire que c'eft la méme Plante. On la nomme au Sud de la Caro-
line Caffena: @ /a Virginie, & au Nord de la Caroline elle eft
connue fous le nom d’Yapon; & dans le dernier de ces endroits elle
cf autant en ufage parmi les Blancs gue parmi les babitans, dis
moins ceux qui demeurent fur la céte.
es
Ilex vomitoria and, as such, must be taken into account.
At first glance, it would seem that Cassine Paragua
Mill. might properly be the basis of a new combination
under J/exv. Closer examination will reveal, however,
that its status is that of a homonym with an orthographic
variation and is, therefore, unavailable.
In the 9th edition of Miller’s ‘‘Gardener’s Dictionary’
(1797) Cassine No. 2, the orthography is altered from
Cassine Paragua to conform with Linnaeus’ C. Peragua.
Thomas Martyn, editor of the work, reported under
Cassine No. 2, that: ‘‘Cassine Paragua of Miller is Ilex
vomitoria. C. Peragua of the Mantissa, is Viburnum
laevigatum.’*> Martyn, having corrected Miller’s ortho-
graphic error, united two names for the Old World con-
cept correctly under the older name Cassine Peragua.
But he then proceeded to misapply this name to the
New World concept which we now know as Ilex vomi-
toria which is ‘‘native of Virginia and Carolina.’’ In con-
sideration of these circumstances, then, Miller's specific
epithet Paragua cannot be available for transfer to Zea.
The confused use of the name I/ex Cassine, which has
likewise hampered our understanding of J/ea vomitoria,
is attributable to the unfortunate fact that this binomial
has been published by two authors for different concepts.
The earlier lev Cassine Linnaeus (Sp. Pl. (1753)
125) was an unfortunately chosen name, for while the bi-
nomial itself refers without any doubt to the dahoon of
the southeastern United States, the specific epithet was
taken from one of the common names of the yaupon
holly and applied to the other holly by Linnaeus in er-
ror. Ilex Cassine L., therefore, is the accepted technical
name for the dahoon holly.
A few years later, Walter (FI. Carol. (1788) 241) pub-
lished Ilew Cassine to refer to the concept we now call
I. vomitoria, as ascertained by reference to a photograph
[ 103 ]
of the type specimen (Schubert [photographer]: ‘* The
Herbarium of Thomas Walter at the British Museum
(Natural History), South Kensington, London, Eng-
land,’ (1946-7) I/eaw Cassine [a bound volume of photo-
graphs in the Gray Herbarium]). Although Walter’s
specific epithet is correctly applied to the plant which
is widely known by that common name, and although
it antedates Ilex vomitoria by one year, the name falls
because it is a homonym.
This dual use of the same binomial has, as Loesener (in
Bot. Centralbl. 47 (1891) 161-168) pointed out, led to
contusion, although even arapid examination of the case
would have made clear all points of uncertainty. E. M.
Hale’s very excellent historical treatment of the black-
drink plant (‘‘Z/ew Cassine, the aboriginal North Ameri-
can tea’’: Bull. No. 14, U.S.D.A. Div. Bot. (1891)), for
example, suffers from his use of the name Ilea Cassine
without even one reference to indicate whether the bi-
nomial of Linnaeus or that of Walter is being accepted ;
Hartwich’s monumental work on narcotic plants (‘‘Die
Menschlichen Genussmittel’’ (1911) 468) uses [lea Cas-
sine L.: and Wehmer (‘‘Die Pflanzenstoffe’’ (1911) 456 ;
[ed. 2] 2 (1981) 718) accepts I. Cassine Walt. The first
chemical report (Smith, H. M.: in Am. Journ. Pharm.
44 (1872) 216) of the discovery of caffeine in a North
American Jlex is of doubtful value partly because of an
uncertainty as to which Ilew Cassine was being examined.
In 1888, still another report of a chemical analysis (Ven-
able, F. R.: in Chem. News 52 (1885) 172) employed
the name J/ex Cassine without author citation, thus leav-
ing room for uncertainty as to the exact species which
was analysed. An even more interesting result of the
confusion of the L/ew Cassine of Linnaeus with that of
Walter is found in a standard pharmacological work
(Tschirch, A.: ‘‘Handbuch der Pharmacognosie”’ 38,
[ 104 J
Abt. 1 (1928) 482), in which its writer ‘‘affirms and denies
the existence of caffeine in I/ex Cassine all in the same
paragraph’’ (Maxwell, M. M.: ‘‘Tlea Cassine L. ... 7’
ms. report (no date) in Libr. Econ. Bot. Harvard Univ. ).
Loesener (in Bot. Centralbl. 47 (1891) 168) rejects Ilex
vomitoria in favor of Ilex caroliniana (Lam.) Loes., a
combination which he made on the basis of Lamarck’s
Cassine caroliniana. This procedure is not permissible.
Lamarck’s binomial Cassine caroliniana (1788) is not
available, since it is a substitute for the validly published
Cassine corymbosa Miller (Dict., [ed. 8] (1768) Cassine
No. 1; [cones (1760) 88, t. 1). It might be pointed out
in passing that Miller’s Cassine corymbosa does not reter
to the concept Ilex vomitoria.
The other synonyms of J/ew vomitoria have not, for
the most part, entered into the chemical, pharmacologi-
‘al and anthropological literature. They are included in
the foregoing enumeration for the sake of completeness.
Several of the Rafinesque names are referred to Ilex
vomitoria with some reserve. They are not based on
specimens which might be examined, and the descrip-
tions are brief and vague in the extreme. It is fortunate
that they do not need to be considered nomenclatorially
in determining the correct name of the yaupon, but, be-
ing later, can be relegated immediately to synonymy.
[ 105 |
.o
A NEW VARIETY OF ILEX VOMITORIA
FROM SOUTHERN MEXICO’
BY
A. J. SHARP
Ilex vomitoria [ Soland. in] Aiton var. chiapensis
A. J. Sharp var. nov.
Haec variatas ab Ice vomitoria principaliter foliis pu-
bescentibus et ramulorum villis longioribus differt.
From the species, this new variety differs primarily in
having loosely pubescent leaves and longer (about twice
as long) hairs on the branchlets. There is a tendency for
the leaves to be a little more ovate and obtuse at the
apices, although many leaves match in shape those from
North American material of the species. The material
of Ilex vomitoria in the Gray Herbarium (all from south-
eastern United States) has been examined, and the leaves
are glabrous, except for occasional puberulence on the
mid-rib at the base.
This report adds another species to the growing list
of plants which occur both in central or southern Mexico
and in southeastern United States. In Mexico, their
ranges are disjunct from their areas in the United States,
and the largest number of such species seems to be found
in the highlands toward the Gulf of Mexico. <A fuller
discussion of the phytogeographical problem basic to
this interrupted distribution will be presented later.
‘Contribution from the Botanical Laboratory, The University of
Tennessee, N.S. 118.
[ 107 ]
Mexico: Estado de Chiapas, south side of Rio Tzaconeja near San
Gregorio. Alt. 6500 ft. November 20, 1945, Aaron J. Sharp & Efrain
Hernandez Xolocotzi 45989. (Tyre in Herb. Gray; Dupticatre Typrs
in Herb. Univ. Tenn., Herb. Inst. Biol. (México) ).—Estado de Chi-
apas, near Lake Jusnajav, about 16 km. northeast of Comitan, Alt.
5500 ft. April 29, 1945, daron J. Sharp & Edward J. Alexander 45444.
—Kstado de Chiapas, moist slope below Morelia, toward Rio Tzacon-
eja, on the trail to San Carlos. Alt. 4600 ft. April 10, 1946, Kfrain
Hernandez Xolocotzi & Aaron J, Sharp 46249.
[ 108 |
BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY
VoL. 14, No. 5
CamsBripGar, Massacnusetts, Marcu 20, 1950
PLANTAE AUSTRO-AMERICANAE VI
PLANTARUM PRINCIPALITER VALLIS AMAZONIAE
NOVARUM VEL MINUS COGNITARUM
NOTAE DIVERSAE
BY
RicuHarp Evans ScHULTES'’
REcENT work on a number of South American plant
collections, for the most part of Amazonian origin, has
uncovered several novelties and has provided us with
additional information of a phytogeographical or ethno-
botanical nature. The following notes are offered as a
contribution to our knowledge of this interesting part
of the New World.
I am grateful to a number of my botanical colleagues
for their determination of plants in certain groups: Dr.
L. H. Bailey and Dr. H. Emery Moore (Palmae); Dr.
Robert Woodson (Apocynaceae); Dr. José Cuatrecasas
(Chloranthaceae, Moraceae, and Musaceae); Dr. Harold
N. Moldenke (Menispermaceae); and Dr. Bassett Ma-
guire (Rapateaceae). It is also a pleasure to express my
appreciation to Mr. Elmer W. Smith for his beautiful
and highly accurate drawings of several of the newly de-
scribed species. The drawings of the concepts of Her-
rania were made possible by a grant from the American
'Botanist, Bureau of Plant Industry, Soils, and Agricultural Engi-
neering, Agricultural Research Administration, United States Depart-
ment of Agriculture; Research Fellow, Botanical Museum, Harvard
University.
[ 109 ]
Cocoa Research Institute, for which my sincere thanks
are due.
TRIURIDACEAE
Soridium Spruceanum Miers in Proc. Linn. Soc.
2 (1850) 75.
Although rare in collections, Soridiwm Spruceanum,
the type of which was from Para, is not uncommon in
the upper Rio Negro basin.
Brazi_: Estado do Amazonas, Rio Uaupés, Taracué. January 28-
February 9, 1948, Richard Evans Schultes & Francisco Lépez 96814.
PALMAE
Bactris balanophora Spruce in Journ. Linn. Soc.
11 (1871) 146, 153.
Bactris balanophora was described from material gath-
ered by Spruce from San Carlos on the upper Rio Negro
in Venezuela. The collection cited below places the con-
cept for the first time well within the Amazonian area
of Colombia and very greatly extends the known range
of the plant.
Cotompia: Comisaria del Amazonas, Rio Igaraparand, vicinity of
La Chorrera. ‘‘Leaf and petiole 9 feet long. Plant 12-15 feet tall.’’
June 4-10, 1942, Richard Evans Schultes 3938.
Chamaedorea integrifolia (7Z'rai/) Dammer in
Verhandl. Bot. Ver. Brandenburg 48 (1907) 125.
This small palm is apparently new to the flora of Co-
lombia.
Co.toms1A: Comisaria del Amazonas, Rio Loretoyacu. Alt. about
100 m. October 20-30, 1945, Richard Evans Schultes 6626.
Chamaedorea lanceolata (Ruiz & Pav.) Kunth
Enum. PI. 3 (1841) 172.
Chamaedorea lanceolata is herewith reported, appar-
ently for the first time, from Colombia.
[ 110 |
Cotompra: Comisaria del Amazonas, Rio Loretoyacu. Alt. about
100 m. ‘‘Fruit ripening orange.’’ October 20-30, 1945, Richard
Evans Schultes 6603.
Geonoma interrupta (wiz & Pav.) Martius Hist.
Nat. Palm. 2 (1823) 8, t. 7.
According to Dugand’s enumeration of palms found
in Colombia (in Caldasia 1 (1940) 44-48), this species of
Geonoma has never before been recorded.
CoLoms1A: Comisaria del Amazonas, Rio Loretoyacu. Alt. about
100 m. ‘“Twelve feet tall.’’ October 20-30, 1945, Richard Evans
Schultes 6749.
Geonoma laxiflora Martius Hist. Nat. Palm. 2
(1823) 12, t. 11.
Geonoma laxiflora, an Amazonian species, is now re-
corded as an element of the flora of Colombia.
Cotomstra : Comisaria del Amazonas, Rio Loretoyacu. Alt. about 100
m. ‘‘Small palm. In clumps, 12-15 ft. tall. Tikuna name = be-ee-chd.”?
October 20-30, 1945, Richard Evans Schultes 6656.
Geonoma pycnostachys Martius Hist. Nat. Palm.
2 (1823) 16, t. 17.
Described from material collected by Martius in the
Caqueta of Colombia, Geonoma pycnostachys is now
registered from the headwaters of the Rio Putumayo.
Since the locality cited below is exactly on the bound-
ary between Colombia and Ecuador, this species should
likewise be assigned to the flora of Ecuador.
Cotoms1a: Comisaria del Putumayo, Rio San Miguel or Sucumbios,
Santa Rosa and vicinity. ‘‘Small terrestrial plant. Kofan Indian name:
tsau-hess.’’ April 1942, Richard Evans Schultes 3547.
Lepidocaryum tenue Martius Hist. Nat. Palm.
2 (1823) 51, t. 47.
Dugand (loc. cit. 31) enumerates only two species of
Lepidocaryum from Colombia. Lepidocaryum tenue, fre-
eile
quent in Amazonian Brazil, is, however, now definitely
reported from Colombia.
Cotomsia: Comisaria del Amazonas, Rio Karaparand, between El
Encanto and the mouth. Alt. about 150 m.? ‘‘Stems used in basket-
ry.’’ May 22-28, 1942, Richard Evans Schultes 3851.
Parascheelia anchistropetala Dugand in Cald-
asia 1 (1940) 12, t. 4, 5.
Schultes 5856 is topotypical and represents the second
collection of this interesting savanna palm.
CotompiA: Comisaria del Vaupés, Rio Vaupés, savanna at base of
Cerro Circasia. ““Inflorescence basal. Leaf 6-7 feet long with 72’
leaflets.’’ March 7, 1944, Richard Evans Schultes 5856.
RaAaPATEACEAE
Rapatea longipes Spruce ev Kérnicke in Linnaea
87 (1871-72) 472.
Spruce collected the type of Rapatea longipes at ‘‘Pa-
nuré’’ on the Rio Uaupés. The collection cited below
is topotypical.
Brazit: Estado do Amazonas, Rio Uaupés, Ipanoré. ‘‘Bracts of in-
florescence red.’’ November 14-15, 1947, Richard Evans Schultes &
Joao Murga Pires 9079.
Rapatea modesta Maguire sp. nov.
Herbae parvae acaulescentes perennantes. Folia 3-6,
20-50 cm. longa; vaginis valde equitantibus distichisque ;
in uno plano dispositis, submembranaceis, vel tenuiter
chartaceis, late rotundatis, nervis distinctis ca. 40 muni-
tis, 1.8-2.5 cm. e costa ad marginem, (4—) 5—9 em. lon-
gis; petiolis (7—) 10-80 cm. longis, ca. 2.4 mm. crassis,
apicem versus anguste alatis; laminis tenuiter chartaceis,
8-15 cm. longis, 8-5 cm. latis, oblongo-ellipticis, nervis
primariis 15-20 (—80), apice subconduplicato, abrupte
caudato-acuminato, obtuso, 15-20 mm. longo, — basi
abrupte constricta, inaequilaterali. Pedunculus pseudo-
[112 ]
terminalis, solitarius, 10-30 em. longus, 2-2.5 mm. cras-
sus, ad apicem paullum dilatatus compressusque, api-
cem versus castaneo-furfuraceus, demum glabrescens.
Inflorescentia compressa; involucri bracteis 2, reniformi-
‘raudatis, 2.5-8.5 cm. longis, 2.5-8 em. latis, tenuiter
multinervatis, albidis, ad basim castaneo-furfuraceis,
apice 1-2.5 ecm. longo; receptaculo convexo, ca. 6 mm.
lato, 4 mm. alto, bracteis paleaceis; spiculis sessilibus,
bracteolis 8-10, lineari-oblanceolatis, 9-10 mm. longis,
1—2 mm. latis, trinervatis, membranaceis, sub calyce ap-
proximatis, abrupte acuminatis, acumine 1.5—2 mm.
longo. Sepala 8, ca. 10 mm. longa, indistinete 3-5 ner-
vata, ad basim delicate membranacea in tubum connata,
sursum libera lanceolato-acuminata, indurata; corolla
delicate membranacea, alba ut videtur, ca. 10 em. longa,
ad basim per 1—2 mm. ad corollam adnata, laminis ovato-
deltoideis, acutis, ca. 5 mm. longis; staminibus 6, ca. 6
mm. longis; filamentis membranaceis, 1-2 mm. longis,
0.8-1 mm. latis; antheris ca. 4 mm. longis, quadriloba-
tis in sectione transversali, ad basim auriculatis sursum
arctantibus, albis ad apicem vix castaneo-furfuraceis; ap-
pendice ca. 1.5 mm. longa, castanea, cochleariformi,
lineari-oblonga, e basi brevi sigmoidea oriente; stylo
plusminusve stamina aequante, stigmate trigono, acuto;
ovario triloculari, ad basim membranaceo, sursum sub-
indurato, albo; ovulum unum in loculo; semine oblongo,
5-6 mm. longo, 3-3.5 mm. lato, integumento interiore
denso, saturate castaneo, longitudinaliter striato, striis
manifestis per integumentum exterius albidum translu-
cens ad apicem disjunctum et bulbosum, ad hilum den-
sum et oblique concavum.
Small acaulescent perennial herbs. Leaves 8 to 6, 20-50
em. long; sheaths strongly equitant and distichous, dis-
posed in a single plane, sub-membranaceous or thinly
chartaceous, broadly rounded, with about 40 distinct
[ 118 ]
nerves, 1.8-2.5 cm. broad from midrib to margin, (4)
5-9 cm. long; petioles (7) 10-80 cm. long, about 2.5
mm. thick, narrowly winged toward the apex; blades
thinly chartaceous, 8-15 cm. long, 3-5 cm. broad, ob-
long-elliptic, with 15-25 (80) primary veins, the apex
sub-conduplicate, abruptly caudate-acuminate, obtuse,
15-20 mm. long, the base abruptly constricted, inequa-
lateral. Peduncle pseudo-terminal, solitary, 10-30 cm.
long, 2-2.5 mm. thick, somewhat enlarged and com-
pressed at the summit, brown-scurfy-pubescent upward,
at length glabrescent. Inflorescence compressed. Invo-
lucral bracts 2, reniform-caudate, 2.5-8.5 cm. long, 2.5-38
cm. broad, finely multi-veined, whitish and brown-scurfy
at the base, the apex 1—2.5 cm. long. Receptacle con-
vex, about 6 mm. broad, 4 mm. high, bracts paleaceous.
Spikelets essentially sessile, bractlets 8-10, linear-oblan-
ceolate, 9-10 mm. long, 1-2 mm. broad, 8-nerved, mem-
branaceous, abruptly acuminate, the acumen 1.5—-2 mm.
long, the bractlets approximately arranged below the
calyx. Sepals 8, about 10 mm. long, faintly 3—-5-veined,
the bases delicately membranous and completely connate
into a tube, the upper halves free, lanceolate-acuminate,
indurate. Corolla delicately membranous, white, appar-
ently about 10 cm. long, the claws connate for 1-2 mm.,
the blade ovate-deltoid, acute, about 5 mm. long. Sta-
mens 6, about 6 mm. long; filaments membranous, 1—
2 mm. long, 0.8-1 mm. broad, adnate to the corolla
for 1-2 mm. ; anthers about 4 mm. long, 4-lobed in cross
section, auriculate at the base tapering upward, white,
scantily brown-scurfy at the summit; appendage about
1.5 mm. long, brown, cochleariform, linear-oblong, at-
tached by ashort, sigmoid base. Style more or less equal-
ling the stamens, the stigma trigonous, acute. Ovary
3-celled, membranous at the base, the upper part subin-
durate, white: ovules 1 per cell. Seed oblong, 5-6 mm.
[114]
long, 8-8.5 mm. broad, the inner integument dense, dark
brown, longitudinally striate, the striae visible through
the whitish translucent outer integument which becomes
separated and bulbous at the apex, and dense and ob-
liquely concave at the hilum.
Doctor Maguire writes me, in connection with his de-
scription of Rapatea modesta, that: ‘‘ Rapatea modesta
is most closely related to PR. /ongipes Spruce ex Kérnicke,
from which it differs in its smaller habit, broader, shorter,
and more conspicuously veined membranous leaf-sheaths,
relatively broader and more numerously veined leaf-
blades, considerably larger heads, more acuminate spiked
bractlets, and white rather than yellow flowers.
‘’These two species are to be associated in habitat and
range. Rapatea modesta occurs in sand savannahs in the
trapécio region between the Putumayo and Amazon
watersheds in southeastern Colombia. The four collec-
tions of Rapatea longipes which I have examined are
from the region of the upper Rio Negro in Brazil, Co-
lombia and Venezuela.
‘The epithet modesta was suggested by the collector’s
comment that this attractive and distinct little Rapatea
appealed to him in the field as a shy, almost retiring,
but lovely member of the caatinga community.’
Cotomstia: Comisaria del Amazonas, Trapécio amaz6nico, interior
regions of trapécio between the Amazon and Putumayo watersheds.
Alt. above 100 m. November 1945, Richard Evans Schultes 6900 (Typr
in N.Y. Bot. Gard.).—Same locality, November 1946, George A.
Black & Richard Evans Schultes 46-361.
Rapatea modesta has a red-flowered form which grows
promiscuously with the white color-form. Red flowers
have not hitherto been reported for the genus.
Cotomsi1A: Comisaria del Amazonas, Trapécio amazonica, interior
regions of trapécio between the Amazon and Putumayo watersheds.
Alt. above 100 m. “Same as 6900, except flowers red.” November
1945, Richard Evans Schultes 6899,
[ 115 ]
Rapatea Spruceana Kérnicke in Linnaea 87 (1871-
73) 470.
Rapatea Spruceana has apparently not hitherto been
reported for the flora of Colombia. The type was col-
lected by Spruce along the lower Rio Guainia, possibly
on the Colombian side. Schultes & Lopez 9297 may be
considered as topotypical.
CotomBia: Comisaria del Vaupés, Rio Negro, El Castillo, San Felipe
(opposite San Carlos, Venezuela). “‘Flowers yellow.’? December 12,
1947, Richard Evans Schultes & Francisco Lopez 9297.
MUSACEAE
Heliconia lingulata Ruiz & Pavon FI. Peruv. et
Chil. 8 (1802) 71.
Known from the Departamento del Valle in Colombia,
Heliconia lingulata, through the collection here cited,
is now recorded from the Colombian Amazon watershed.
Cotompia: Departamento del Cauca, Puerto Limon, ‘‘Bracts rose-
coloured. Platanillo, Leaves 12 feet long. Inflorescence 6 feet long.”’
February 28, 1942, Richard Evans Schultes 3351.
Heliconia psittacorum Linnaeus filius Suppl.
(1781) 158.
Heliconia psittacorum is widespread in the northern
tropical parts of South America; in Colombia, it has
been collected from the Llanos, from Santander, and
from the Vaupés basin. Schultes 6633 is apparently the
second collection from the Colombian Amazonia and
greatly extends the known range of the species in that
area.
Cotomp1a: Comisaria del Amazonas, Rio Loretoyacu. “*Bracts red
with tips green. Capavira.’’ October 20-30, 1945, Richard Evans
Schultes 6633.
Heliconia Schumanniana Loesener in Engl. Bot.
Jahrb. 54, Beibl. 117 (1916) 12.
Heliconia Schumanniana, the type of which came from
[ 116 |
eastern Pert, has apparently not previously been re-
corded from Colombia.
Cotompia: Comisaria del Putumayo, Rio Caucaya, entre Puerto
Jaramillo y el Rio Putumayo. Altitude about 225 m. May 16, 1942,
Richard Evans Schultes 3711.
Heliconia stricta Huber in Bol. Mus. Goeldi 4
(1906) 543.
Heliconia stricta was described from material collected
in Pert. Schultes 3489, from the boundary of Colombia
and Ecuador, is apparently the first collection of the spe-
cies from either of these countries.
Cotoms1a: Comisaria del Putumayo, Rio Sucumbios (San Miguel)
Puerto Conejo. “‘Platanillo.’’ April 2-5, 1942, Richard Evans Schultes
3489.
CHLORANTHACEAE
Hedyosmum toxicum Cuwatrecasas in Caldasia 8
(1945) 432.
Hedyosmum toxicum is very seldom used as a medicine
and always with great care, and it is never taken as a
beverage. Unless the decoction of the leaves be made
rather weak, it can act as a violent emetic and is a strong
intoxicant.
CotomspiA: Comisaria del Putumayo, Paramo de San Antonio, alti-
tude about 3000 m. “‘Granisillo pequeto. Large tree. Leaves aromatic.
A tea of leaves sometimes used as a stomach tonic and as purge. It
is a strong emetic.’’ February 13, 1942, Richard Evans Schultes 3249.
Hedyosmum translucidum Cuwatrecasas in Cald-
asia 8 (1945) 436.
A remarkable tonic-stimulant effect is noted immedi-
ately after a draught of a very hot decoction of the leaves
of Hedyosmum translucidum. It may contain an active
stimulant. During plant-collecting trips in the high, cold
piramos of the western part of the Comisaria del Putu-
mayo, I often prepared from the leaves of this bush,
granisillo, the delicious aromatic beverage which travel-
[117 ]
lers in these high, inhospitable regions have learned to
employ as a stimulant.
Cotompra: Comisaria del Putumayo, Péramo de San Antonio, alti-
tude about 3000 m. “‘Granisillo del grande. Leaves aromatic. Tea of
leaves used occasionally as a beverage and medicinally for stomach
upsets.’” February 13, 1942, Richard Evans Schultes 3221.
MoRACEAE
Cecropia latiloba Miquel in Martius FI. Bras. 4,
pt. 1 (1858) 147.
Known from Brazil and possibly Pert, Cecropia lati-
loba seems not to have been previously reported as a
component of the Colombian flora.
Cotomsra: Comisaria del Amazonas, Rio Loretoyacu. ““Small tree.”’
October 20-30, 1945, Richard Evans Schultes 6720.—Same locality.
**Small tree.’? November 1945, Richard Evans Schultes 6912.
Cecropia telealbida Cuatrecasas in Rev. Acad.
Colomb. Ciéne. 6 (1945) 294, t. 2, fig. 6.
The type of Cecropia telealbida was collected in south-
ern Huila at a much higher altitude (1850 m.) than the
two collections cited below.
Cotomsia: Departamento del Huila, Rio Villalobos, Quebrada Guay-
abo, altitude 1400-1450 m. January 1943, Richard Evans Schultes &
M. Villarreal 5172.—Departamento del Huila, Rio Villalobos, region
of the confluence of Rios Villalobos and Cauchos. Altitude 1400 m.
January 1943, Richard Evans Schultes & M. Villarreal 5205.
Cecropia telenivea Cuatrecasas in Rev. Acad. Co-
lomb. Ciéne. 6 (1945) 295.
This second collection of Cecropia telenivea is topo-
typical.
Cotomsia: Comisaria del Putumayo, Valle de Sibundoy, Sibundoy,
hills north of town, altitude about 2250 m. ‘‘Large tree.’’ February
18, 1942, Richard Evans Schultes 3276.
Cecropia tolimensis Cuatrecasas in Rev. Acad.
Colomb. Ciéne. 6 (1945) 282, t. 1, fig. 4, t. 2, fig. 5.
[ 118 ]
Cecropia tolimensis was described from material from
Tolima, much to the north of the Huila locality of
Schultes & Villarreal 5128 which represents the second
collection of the species to be reported.
Cotoms1A: Departamento del Huila, Pitalito, Calamo, open fields.
Altitude about 1300 m. December 30, 1942, Richard Evans Schultes
& M. Villarreal 5128.
Coussapoa intermedia Martius ex Miquel in Mar-
tius Fl. Bras. 4, pt. 1 (1858) 133.
Coussapoa intermedia, hitherto known from Amazon-
ian Pert and Brazil, is now recorded for the flora of
Colombia.
Cotoms1a: Comisaria del Amazonas, Rio Loretoyacu. ‘‘Large tree.’’
October 20-30, 1945, Richard Evans Schultes 6726.
Coussapoa magnifolia 7?écu/ in Ann. Sci. Nat.,
ser. 3, 8 (1847) 98.
Although Coussapoa magnifolia is known from eastern
Pert, I have been unable to find other collections of
this species from Colombia.
Cotoms1a: Comisaria del Amazonas, Rio Loretoyacu. ‘“Epiphyte.’’
October 20-30, 1949, Richard Evans Schultes 6693.
OLACACEAE
Heisteria cyanocarpa Poeppig & Endlicher Nov.
Gen. ac Sp. 3 (1845) 35, t. 241.
In Macbride’s ‘‘Flora of Peru’’ (Field Mus. Publ.
Bot. 13, pt. 2, no. 2 (1987) 423), Standley gives, as the
distribution of this species, eastern Perti and Amazonian
Brazil. Apparently it has not been recorded as an ele-
ment of the Colombian flora, although it is frequent in
the southern part of the Colombian Amazonia. Several
of the collections cited as Peruvian, however, are actually
from Colombian territory and attest the frequency of
this shrub, especially in the trapécio amazdénico. Ll.
pice
Wilhams 2745, 2805, 2808, 2949 (from La Victoria) and
3169 (from Leticia) are Colombian.
Similarly, Heisterta eurycarpa Standl., known only
from the type collection made at La Victoria, near Leti-
cia, Colombia, has been cited as Peruvian; although this
concept undoubtedly occurs on the Peruvian as well as
on the Colombian side of the Amazon River, it is advis-
able to include it in an enumeration of Colombian plants.
Cotoms1A: Comisaria del Amazonas. Trapécio amazénico, Loreto-
yacu River. Alt. about 100 m. ‘‘Bush. Bracts red.’? October 1945,
Richard Evans Schultes 6832.
MENISPERMACEAE
Sciadotenia toxifera Krukoff & Smith in Bull.
Torr. Bot. Club 66 (1989) 808.
Sciadotenia towifera has previously been known from
Amazonian Ecuador where Richard C. Gill collected it
as an ingredient of the curare of the Canelos Indians of
the Napo-Pastaza area. This was the first report of the
genus as an arrow poison plant (Krukoff & Smith loe.
cit.).
The Witoto Indians of Amazonian Colombia no
longer prepare curare, but knowledge of which plants
entered into their poisons is still extant. Sczadotenia
toxvifera was indicated as one of the principal ingredients.
CotomsiaA: Comisaria del Amazonas, Karaparana, El Encanto.
**Bush in swamps. Fruit with soft brown hairs but with a blue-green
sheen through indumentum. Height 8 ft. Witoto name: he-de-kd-pe.
Formerly used to make arrow poisons, together with other plants.’’
May 23, 1942, Richard Evans Schultes 3866.
STERCULIACEAE
Herrania Camargoana R. E. Schultes sp. nov.
Arbuscula parva, tenuis gracilisque, vulgo plusminusve
octo vel decem (sed saepe usque ad viginti septem) pedes
alta; erecto cum trunco tereti, circiter 4-5 pollices in
[ 120 ]
diametro, nigro cum cortice obtecta; prope trunci api-
cem parce ramosa; ramis tomentosis sed mox glabratis.
Ramuli dense villosi, ferrugineis cum pilis, subglabres-
centes. Folia amplissima, digitata, sex- ad novem-foliata,
longissime petiolata. Petioli teretes, basi valde constricti,
molliter aureo-ferrugineo-tomentelli, usque ad 60 ecm.
longi, 10 mm. in diametro. Stipulae persistentes, subu-
latae, densissime tomentellae, usque ad 8 cm. longae.
Foliola sessilia, oblanceolata vel late lanceolato-ovata,
leviter erecta, inaequalia, membranaceo-papyracea, apice
acuminata, basi attenuata, margine dimidio superiore
regulariter et conspicue sinuata, et omnino pilis vel pseu-
dociliis stellatis armata, 60-75 cm. longa, 16-26 em. lata,
supra aspera, sparse pilosa cum pilis longis et solitariis,
subtus submolliter tomentella cum pilis longis stellatis.
Inflorescentiae fasciculatae, multiflorae, ex trunci omni-
bus partibus sed vulgo inferioribus prorumpentes. Pedi-
celli articulati, usque ad 28 mm. longi, 0.8 mm. in
diametro. Alabaster floris globosus, usque ad 10 mm. in
diametro, stellato-pilosus. Calyx trifidus, fere usque ad
basim divisus, subeymbiformis. Sepala late elliptico-
oblonga, apice subacuta, margine integra, extus atropur-
purea, intus sanguinea, plusminusve 12 mm. longa, 8-9
mm. lata, intus glabra, extus stellato-pilosa cum pilis
ferrugineis usque ad 1 mm. longis atque pilis albis mi-
nutissimis. Petala quinque, sessilia, obovato-rotundata,
apice valdissime concavo-cucullata, circiter 8 mm. longa,
6 mm. lata, quinque cum nervis atropurpureis longitu-
dinalibus atque nervulis reticulatis, alibi flava, extus con-
spicue muricato-verrucosa, superne in ligulam extensa.
Ligulae lineares, plusminusve 90 mm. longae, basi 1.7
mm. latae, apice filiformes, basi ipsa sanguineae sed max-
ima pro parte albido-flavae. Tubus stamineus quinque-
divisus cum staminibus diantheriferis et simplicibus cum
filamentis brevibus liberisque. Ovarium ellipsoideum,
[121]
3.5 mm. longum, 2-2.5 mm. in diametro, densissime et
grossiuscule albido-pilosum. Stylus teres, simplex, flavus,
stigmate apice inconspicue quinquediviso, 3 mm. longus.
Staminodia conspicua, rhomboideo-elliptica, apice obtu-
sa, Margine integra, utrinque verrucosa, cinereo-purpurea,
14 mm. longa, 5 mm. lata. Fructus numerosi, quasi glo-
bosi vel leviter ellipsoidei, apice abruptissime et longe
apiculati (apicula 2-2.5 cm. longa), plusminusve 8-8.5
em. longi, 8.5-4.5 cm. in diametro, basi attenuati, sepa-
lis persistentibus cum pedunculo 3-4 cm. longo, 2.5 mm.
in diametro, longitudinaliter decem-costati, cum costis
primariis et secundariis subaequalibus, tenuibus cultri-
formibusque, altitudine irregularibus sed maxime pro
parte (vivo) 5-6 mm. altis, transverse irregulariter sed
conspicue costati cum costis cultriformibus, quam longi-
tudinalibus paulo humilioribus vel saepe altioribus, in
costarum longitudinalium et transversalium junctionibus
projectionibus carnoso-mollibus mammoso-spiniformibus
(quae apice aliquid hebetatae sunt) productis, in costis
omnibus et multo sparsius inter costas pilis stellatis urti-
cantibus armati; pericarpio crasso, maturitate rufo vel
sanguineo; semina viginti quinque, obtuse rotundato-
pyramidalia, circiter 9 mm. longaX11 mm. lataX7 mm.
crassa, in pulpa alba inclusa.
Brazit: Estado do Amazonas, Rio Negro, Serra de Sao Gabriel.
‘*Treelet 12 feet tall. Fruit globose. All ten longitudinal ribs about
equal, sharp, transverse ribs as high or higher. Projections formed at
juncture of longitudinal and transverse ribs long, upturned, soft, his-
pid, stinging hairs along ribs and to a lesser extent between. Ripen-
ing scarlet. Seeds 25.’’ February 12, 1948, Richard Evans Schultes &
Francisco Lépez 9722 (Tyrer in Herb. Gray ),—Estado do Amazonas,
Rio Uaupés, between Ipanoré and confluence with Rio Negro, Serra
Wabeesee, on left bank a little below Bela Vista. November 17, 1947,
Richard Evans Schultes & Francisco Lépez 9144 (Tyrer of flowers in
Herb. Gray ).—Estado do Amazonas, Rio Negro. ‘‘Encosta da Serra
de Cabary a 450 m. de altitude. Arbusto de 5 metros.’* December 2,
1945, Ricardo de Lemos Frées 21468.—Fstado do Amazonas, Rio Ne-
gro, Uaupés (Sao Gabriel). ‘“Mata sopé Serra Sao Gabriel. Arbusto
[ 122 ]
5 metros; flores ruges rubro.’” December 22, 1945, Ricardo de Lemos
Fro6es 21540.—Estado do Amazonas, Rio Negro, Sao Gabriel (Uaupés),
Serra de Sao Gabriel, near summit. Alt. c. 100 m. ‘*Single-trunked
treelet up to 9 m. tall. Three or four leaves on top. Fruit clustered
on stem, usually near base, three to four in each fascicle, almost glo-
bose with very abruptly pointed apex, main and secondary longitudi-
nal ribs almost equal, but transverse ribs as high or higher than main
ribs, forming upturned pseudoechinate projections at right angled
point of juncture, hispid stinging hairs all along ribs and to lesser ex-
tent on surface of fruit. Ripening a scarlet. Seeds 25, embedded in
acid, white pulp. Trunk 4—5 inches in diameter. Bark smooth, black.
Leaflets curiously ‘‘spurred’’ at base, thickened spur causing them
to be somewhat erect. Extremely common. Leaflets entire in lower
half, regularly sinuate in upper half. Cacao jacaré. Tukano = ee-sé-pe-
ke.’’ September 15, 1947, Richard Evans Schultes & Francisco Lépez
8758.—Same locality and date. “‘Trunk black, columnar, 3 inches in
diameter. Treelet 8 m. tall. Leaf lobes thickened at base into spur-
like prong, throwing them upwards. Fruit on stem at 5 ft. above
ground. Ripening red. Transverse and main ridges meet to form
curious thickening in form of upturned ‘hooks,’ covered with stinging
hairs.’’ Richard Evans Schultes &§ Francisco Lopez 8759.—Same locality
and date. ‘“Seven m. tall; diameter 3 inches. Bark black. Fruit ripens
red. 25 seeds. Fruit looks softly echinate due to warty projections all
over where transverse and horizontal ribs cross.’’ Richard Evans Schultes
5 Francisco Lopez 8762.—Same locality and date. Richard Evans
Schultes § Francisco Lopez 8763,— Estado do Amazonas, Rio Negro,
Tapurucuara (Santa Isabel). September 11, 1947, Richard Evans Schultes
& Francisco Lopez 8956.—FEstado do Amazonas, Rio Negro basin, Rio
Padauiary. October 27, 1947, Ricardo de Lemos Frées 22673.—FEstado
do Amazonas, Rio Negro, near Sao Gabriel, at base of Serra Uanari.
October 31, 1947, Richard Evans Schultes & Joao Murga Pires 8978 .—
Estado do Amazonas, Rio Uaupés, Serra Wabeesee, below Bela Vista.
**Flowers many in clusters. Calyx segments red, petals and staminodes
greyish purple. Ligules greyish purple, 5-6 cm. long. Treelet 15 ft.
tall.’’ November 17, 1947, Richard Evans Schultes & Joao Murca Pires
9130.—Estado do Amazonas, Rio Negro, Serra de Sao Gabriel. “‘Tree-
let 8 ft. tall. Flowers in many-flowered fascicles. Sepals externally
dark blood-red, internally scarlet; petals ashy red with yellow stripes;
staminodes ashy purple-red, internally showing white-yellow patches,
externally dark red. Ligules 8-9 cm. long, linear, externally red, in-
ternally white-yellow, but folded or inrolled so that red is enclosed.
Tukano: 6-sd-pee-ké.’’ November 20-25, 1947, Richard Evans Schultes
& Francisco Lopez 9162.—Estado do Amazonas, Rio Negro, mouth of
Rio Xié. December 2, 1947, Richard Evans Schultes & Francisco Lopez
9205.—Estado do Amazonas, Rio Negro, Nazaré. ‘‘Treelet 15 ft.
[ 128 |
tall. Trunk 4 inches in diameter. Fascicles with up to $0 flowers.
Flowers dull ash-purple with exterior of sepals scarlet and most of
length of infolded ligules a faintly purplish white. Buds perfectly glo-
bose. Bark nearly black. Leaves few. Leaflets unevenly lanceolate.
Fruit ‘spiny,’ apically very long acuminate. Cacao de jacaré.’’ Decem-
ber 5, 1947, Richard Evans Schultes & Francisco Lopez 9240.—Estado
do Amazonas, Rio Negro, Serra de Sao Gabriel. ‘‘Small treelet, 12
feet tall.’’ January 14, 1948, Richard Evans Schultes & Francisco Lopez
9619.—Estado do Amazonas, Rio Negro, Serra Jacamin. ‘‘Small tree-
let about 12 feet tall. Trunk slender, 3-4 in. in diameter, bark thin,
smooth, black. Leaves large, hairy, upper half of leaflets sinuate. Cacao
de macaco.’’ March 27, 1948, Richard Evans Schultes & Francisco Lopez
9747.— Estado do Amazonas, Rio Negro between Sao Felipe and Kara-
pana. May 4, 1948, Richard Evans Schultes & Francisco Lopez 9869.
Herrania Camargoana is named in honor of Dr.
Felisberto Camargo, founder and director of the Insti-
tuto Agronomico do Norte in Belém do Para, Brazil.
Possessed of a deep patriotism and a fervent desire to
take the benefits of agronomical science to the poor and
forgotten caboclo of the Amazon valley, Camargo has, at
times almost single-handedly, carried out an energetic
crusade to improve the lot of the Amazon through better
agriculture. In spite of discouraging odds in the form
of tremendous distances, primitive transportation and
actual lack of geographical knowledge of large areas;
and of poverty, sickness, apathy and lack of education
amongst the people themselves, the efforts of Camargo
and his small but loyal group of scientists are beginning
to bear real fruit. To the best of my knowledge, this
program, embracing many phases of fundamental biolo-
logical research (from plant exploration and introduction
to forest tree breeding and phytopathology) and the es-
tablishment of nurseries and plantations, constitutes the
only sustained agricultural work of a truly scientific na-
ture which the Amazon has seen in the four centuries
since its discovery by Europeans.
Restricted apparently to the uppermost Rio Negro
basin, Herrania Camargoana seems to have as its closest
[ 124 |
ally the Guianan and Venezuelan HZ. lemniscata. This
relationship is strikingly evident when one compares the
fruits of the two concepts. Both species have relatively
small capsules in which there are transverse ribs nearly
as large as the cultriform longitudinal ribs, and soft,
pointed mammoid projections at each junction of the
longitudinal and transverse ribs. The former species,
however, has much longer and upturned projections than
the latter, and would seem, in this as in some other char-
acters, to represent an extreme in the evolution of the
genus. ‘There would appear to be a rather easily trace-
able trend from Herrania Mariae through Hi. lemniscata
to H. Camargoana, on the one hand, and to A. lacinii-
folia on the other.
The coloration of the flowers of Herrania Camargoana
and FH. lemniscata is similarly complex and also indicates
arelationship. No other known species of Herrania can
match these two for complexity of floral coloration. Her-
rana Camargoana, according to field notes quoted in
detail above, has sepals which are dark blood-red exter-
nally but scarlet internally ; petals which are ashy red or
purple with yellowish stripes; staminodes which are ashy
purple-maroon with white-yellow patches internally but
entirely dark red externally ; and ligules, red without and
white-yellow within, which are folded or inrolled, so that
the red is enclosed and is not seen directly. Herrania
lemniscata has, according to field notes (Steyermark
60558), sepals which are white in the uppermost two-
thirds and rose-salmon below, with rose stripes; and
staminodes (called ‘‘petals’” on the label) which are dull
yellow with dull rose specks in the lower half.
The shape and size of the leaflets, however, differ
strikingly in the two concepts. Herrania Camargoana
has oblanceolate or broadly lanceolate-ovate leaflets
which measure 60-75 cm. in length and 16-26 em. in
Bele
width with the upper half regularly and conspicuously
sinuate. Herrania lemniscata has leaflets which are at
least 80 cm. long and 40 cm. wide with the margin very
deeply pinnatilobate with usually four irregular, for the
most part widely triangular or widely lanceolate-acumi-
nate segments, each up to 18 cm. long and 9-10 cm.
wide.
Herrania kanukuensis FP. I. Schultes in Caldasia
2 (1943) 11; in Bot. Mus. Leafl. Harvard Univ. 13
(1949) 277.
Hitherto known only from the type collection from
the Kanuku Mountains in adjacent British Guiana, er-
rania kanukuensis is now registered, through the collec-
tion cited below, from northern Amazonian Brazil.
Inasmuch as Herrania kanukuensis has never been
illustrated, it has seemed advisable to publish, with this
phytogeographical note, a drawing of this species (Plate
».&.S. EO BY
Brazit: Territorio do Rio Branco. Lower Rio Branco, Tapanaruca.
““Tree in clump, 3 meters; rouge flowers. Lowland, border of high
forest.’’ February 23, 1948, Ricardo de Lemos Frées 23003,
Herrania kofanorum RP. 1. Schultes sp. nov.
Arbuscula parva, tenuis gracilisque, usque ad quinde-
cim pedes alta, ex radice plerumque unus truncus erec-
tus, cylindricus, apice ramosus, cinereo-nigro cum cortice
scrobiculato et scabrido, usque ad septem cm. in diame-
tro crescens. Rami ferrugineo-tomentosi, denique sub-
glabrati, subteretes sulcatique. Ramuli similes sed to-
mentosiores. Folia ad trunci apicem collecta, ampla,
digitata, longissime petiolata, septem-foliolata, stipulata.
Stipulae caducae, lineares, acutae, 2.5-8 cm. longae,
circa 8 mm. latae, siccae, extus hispidulae vel strigillosae,
intus usu subglabrae. Petioli robusti, teretes sed obscuris-
sime suleati, basi leviter dilatati, subferruginei, densissime
[ 126 ]
atque molliter tomentosi, usque ad 80 cm. longi, 6 mm.
in diametro. Foliola sessilia, inaequalia, lanceolato-
oblonga, apice in cuspidem vel mucronem acutum usque
ad 2 cm. longum producta, basi sensim attenuato-decur-
rentia, Margine conspicue atque regulariter undulato-
sinuata; laminis firme coriaceis, plerumque 17-30 cm.
longis, 6-11 cm. latis, supra atroviridibus, glabris vel max-
ime sparsissime atque minutissime strigilloso-pilosiusculis
albis cum pilis caducis, in venis praecipuis minute fusco-
hirsutis, subtus fulvo-viridibus, densissime atque molliter
stellato-pilosis, in venis praecipuis ferrugineo-tomentosis ;
costa nervisque utrinque sed subtus magis prominenter
elevatis. Inflorescentiae fasciculatae, usque ad viginti-
florae. Flores caulini, longe pedicellati, e trunco inferiore
orti, in racemis contractis prorumpentes. Alabaster mag-
nus, elongato-globosus, 1.8 cm. in diametro, stellato-
pilosus, subfulvo-rubens. Pedicelli robusti, teretes, articu-
lati, densissime et minute stellato-pilosiusculi, subcinereis
cum pilis, plerumque 9-10 mm. longi, 1-1.5 mm. in
diametro, basi bractea brevi, lineari, acuta, dense tomen-
tosa, 2-4 mm. longa subtenti. Calyx subcymbiformis,
fere usque ad basim divisus. Sepala tria, valdissime in-
aequalia, subchartacea, margine integra, intus minute
papillosa, subglabra atque probabiliter cito glabra, extus
sparsim pilis stellato-strigosis usque ad 1 mm. longis et
etiam pilis stellato-pilosiusculis minutissimis armata, aes-
tivatione valvata; duo interiora elliptica, apice acuta, 10
mm. longa et 6 mm. lata; exterius rotundato-ovatum,
15-16 mm. longum, 15 mm. latum, apice rotundatum et
saepe profunde (usque ad 2 mm.) incisum, incisura ut
sulcus medianus ad laminae basim intus extensa. Petala
quattuor vel quinque, rotundato-obovata, sessilia, con-
cava, apice valde cucullata, glabra, utrinque (sed extus
magis) muricato-papillosa vel granulosa, atrorubentia,
quinquenervia, purpureis cum venis longitudinaliter stri-
[ 127 ]
ato-venosa, superne in ligulam extensa, 9 mm. longa, 7
mm. lata. Petalorum ligulae filiformes, pendulae, mem-
branaceae, omnino glabrae sed basi minute granulosae,
plusminusve 1 mm. latae, 80-100 mm. longae, basi valde
dilatatae. 'Tubus stamineus quinquedivisus, staminibus
invicem duo- et quattuor-antheriferis, filamentis valde
complanatis, brevibus liberisque. Staminodia conspicue
petaloidea, lanceolato-elliptica, apice acuta, basi attenu-
ata, utrinque grosse muricato-papillosa, margine integra
22 mm. longa, 6 mm. lata. Ovarium sessile, elongato-
ovoideum, distincte decemcostatum atque quinquelocu-
lare, ochraceum, densissime stellato-pilosum, 2.5-8 mm.
in diametro. Pistillum complanatum, 3 mm. longum,
glabrum, simplex. Fructus ignotus sed luteus in maturi-
tate dicitur.
Herrania kofanorum differs from its very close ally H/.
balaénsis Preuss in being smaller, in having leaflets only
half as large, and in having the sepals very conspicuously
unequal instead of nearly alike. Herrania kofanorum has
two inner sepals which are elliptic, 206 mm. and an
outer one which is rotund-ovate, 15-16 X 15 mm., whereas
Hi. balaénsis has three lanceolate-elliptic sepals which
measure 14X6 mm. Furthermore, the outer sepal of
Herrania kofanorum is so constructed that it is often
conspicuously slit to a depth of 2 mm., and this slit is
prolonged as a furrow to the base of the interior of the
sepal (when the split is not present, there is a markedly
thin furrow); nothing similar is seen in H. balaénsis.
Both these species are related to Herrania Dugandu.
Ecvapor: Rio San Miguel o Sucumbios, entre el Rio Putumayo y
la Quebrada Teteyé, alt. 260 m. Nombre Kofan: ko-kee-ot-chu. ‘“Small
tree, 15 ft. Flowers large, deep red-purple in all parts. Fascicles
many-flowered. Cacao silvestre.’’ March 29, 1942, R. E. Schultes 3478.
(Type in Econ. Herb. Oakes Ames; Dupiicatr Types in Herb. Nac.
Colomb., Herb. Gray, U. S. Nat. Herb.).
Cotompt1A: Comisaria del Putumayo, Rio Putumayo, trocha entre
[ 128 ]
Puerto Ospina y Concepcion, alt. 250 m. Nombre vulgar: cacao sil-
vestre, cacao de monte. ‘‘Small tree, 12 ft. tall. Flowers dark red.
Kofan name: ko-kee-ot-chu.’’ April 20-30, 1942, R. E. Schultes 3670.
Herrania Mariae (Mart.) Decaisne ex Goudot var.
putumayonis PF. FE. Schultes var. nov.
Arbuscula usque ad duodecim vel quattuordecim
pedes alta, ab Herrania Mariae principaliter foliolis mul-
to majoribus (usque ad 52 cm. longis, 18 cm. latis),
lanceolato-ellipticis (non conspicue rhomboideis); peti-
olis robustioribus et longioribus; floribus majoribus, ala-
bastro globoso usque ad 17 mm. in diametro; ligulis
brevioribus, usque ad 70 mm. longis sed saepissime paulo
brevioribus differt.
Additional material may indicate that this concept is
deserving of specific rank. At the present time, however,
it would seem advisable to treat it as representing a va-
riety of Herrania Mariae. The fruit of Schultes 4010 is
hardly distinguishable from that of Herrania Mariae.
The flowers have several differentiating characters, the
most conspicuous of which is the shorter ligule. Vege-
tatively, the collection is extremely similar to Herrania
nycterodendron (with the type plant of which it was
growing) and differs markedly from H. Mariae chiefly
in the departure from the typical rhomboid form of the
leaflets and in their unusually large size. The type plant
of Herrania Mariae var. putumayonis consisted of four
or five trunks ina clump, whereas H. Mariae usually, if
not always, is a treelet with a single trunk.
Herrania Mariae var. putwmayonis may represent a
western variant of the species which is most abundant
in the eastern half of the Amazon basin.
Peru: Departamento de Loreto, Rio Putumayo, entre los Rios Iga-
raparand y Yaguas, Puerto San Salvador o Pesqueria, cerca de las Is-
las Batal6n Ly Batalén II. Alt. 100-150 m.(?). ‘‘Bark basally black,
smooth, above grey-brown. Wood soft. Height 12-14 feet. Petals
P4129 |
deep blood-red. Ligules pink, white for half the length. Witoto
name: mu-se-na. Spanish name: cacao silvestre.’? June 20, 1942,
Richard Evans Schultes 4010 (Tyrer in Herb. Gray).
Herrania nitida (Poepp.) R. E. Schultes var. as-
pera (Karsten & Triana) R. EF. Schultes comb. nov.
Brotobroma aspera Karsten & Triana ex Triana Nuev.
jén. e esp. plant. fl. Neo-Granad. (1854) 12.
Herrania aspera (Karsten & Triana) Karsten in Lin-
naea 28 (1857) 447,
Arbuscula parva ab Herrania nitida principaliter foli-
olis utrinque asperioribus et apicem versus margine defi-
nite sinuatis, florum partibus omnibus sanguineo-pur-
pureis, fructibus probabiliter minoribus differt.
With the searcity of collections, it is rather difficult
to evaluate this concept. It is certainly not specifically
distinct from the widespread and somewhat variable
Amazonian Herrania nitida. At the present state of our
knowledge, it is probably best treated as a variety. The
very definitely and regularly sinuate margin of the upper
part of the leaflets is apparently a constant character,
although in several of the Peruvian collections which I
have assigned to Herrania nitida there is a tendency for
the margin to be obscurely undulate.
Karsten, in discussing Herrania aspera (in Linnaea
loc. cit.) and in pointing out its difference from HZ. pul-
cherrima Goudot, stated that it occurs ‘‘in vallis Oreno-
censis marginibus ad pedem Andium bogotensium mer-
idensiumque. ... et littora fluminis Magdalenae.’’ He
cited no specimens. The presumed type of the concept
— Triana 5333—was collected in the Llanos of Villavi-
cencio. The only other collections which have as yet
come to my attention are a topotype, one from the
Apaporis basin and several from the trapécio amazénico.
Therefore, | believe that Karsten’s assertion that Her-
rania nitida var. aspera is found in the Magdalena basin
[ 130 |
may be erroneous and may have been based on a sterile
collection of H. albiflora, superficially often rather similar.
When Triana and Planchon (Prodr. Fl. Novo-Granat.
(1862) 209) erroneously reduced Herrania aspera to syn-
onymy under AZ. pulcherrima, additional confusion con-
cerning the distribution of the concept resulted.
Cotomsia: Intendencia del Meta, Llanos de San Martin, Villavi-
cencio, altitude 400 m., January 1856, J. Triana 5333 (Typer).—Com-
isaria del Amazonas, Rio Amazonas, La Victoria. ‘‘Small tree. Edge
of forest.’’ August-September, 1929, Llewelyn Williams 2816,.—In-
tendencia del Meta, Llanos de San Martin, Villavicencio, altitude
about 500 m. “‘Dense forest. Small tree, 3 m. high. Leaves borne
at summit; stem about 2 ecm. in diameter, bearing red flowers in
lower part.’’ March 17, 1939, EF. P. Killip 34247.—Comisaria del
Vaupés, Upper Apaporis basin, Rio Macaya near confluence of Ajaju
and Macaya, near Puerto Hevea. Altitude 350 meters. Sandy, well-
drained soil. “‘Treelet 12 ft. tall, basal diameter 3 inches. Bark
smooth, black, thin. Fruit on lower portion of trunk, ellipsoid, 10-
ribbed, long-tipped, rich green, stinging hairs up to 1 mm. long along
ribs and between them to a lesser extent. Persistent sepals brown,
dried, densely hirsute. Fruit (without peduncle which is 25 mm. long
and 5 mm. in diameter) 9-10 cm. <5 em, in diameter.’’? June 1943,
Richard Evans Schultes 5529.—Comisaria del Amazonas, Trapécio am-
azonico, Leticia. Alt. about 100 m. September—November 1944,
Richard Evans Schultes 6141; 6143; 6144; 6147; 6149; 6192 A.—Com-
isaria del Amazonas, Trapécio amazénico, Rio Loretoyacu. Alt. about
100 m. “‘Small treelet. Tikuna Indian name: cha-te-rd.’’ October
20-30, 1945, Richard Evans Schultes 6640.
Herrania nitida (Pocpp.) R. KE. Schultes fma.
sphenophylla FR. L. Schultes stat. nov.
Herrania nitida (Poepp.) R. E. Schultes var. spheno-
phylla R. K. Schultes in Caldasia 2, no. 6 (1948) 20.
Continued field studies of the rather variable Herrania
nitida complex indicate the advisability of treating this
concept as a form and not as a variety.
Herrania pulcherrima Goudot var. pacifica PR. HL.
Schultes var. nov.
Arbor parva usque ad viginti quinque pedes alta, ab
[ 131 ]
Herrania pulcherrima foliolis valde membranaceis, supra
minute stellato-pilosis et non muricatis vel subtubercu-
latis, lateralibus saepissime valde obliquis; floribus vulgo
multo minoribus cum partibus (petalis ligulisque) ochra-
ceis vel albis; fructibus minoribus (11.5 cm. longis, 7
em. in diametro) principaliter differt.
This concept would seem to represent a Pacific coastal
variant of Herrania pulcherrima, a species which, in its
typical form, is endemic to the cordilleras of Colombia.
Cotompia: Departamento del Valle, Pacific coast, Rio Yurumangui,
Caimanero. Forest. Alt. 10 m. ‘‘Arbolito 6 m. Tallo 6-7 em. didme-
tro, gris negruzco oscuro, leso con verrugas. Inflorescencias caulinares
paucifloras. Capullos florales pardo-ocraceos.’’ February 5, 1944, J.
Cuatrecasas 16010.—Intendencia del Chocé6, Rio San Juan, vicinity
of Palestina. Alt. 0-30 m. “‘Arbolito 8 m. Tallo 15 em. didmetro.
Hoja membranosa, gruesa, rigida, verde amarillenta. Peciolo verde,
ocraceo. Fruto 11.5 em, <7 em., prismatico-pyramidal, apuntado con
5 costillas muy salientes y otras cortas; algunos nervios muy marca-
dos; verde amarillento.’’ May 28, 1946, J. Cuatrecasas 21337 .—De-
partamento de Antioquia, north of Dabeiba, road to Turbo. “*Closed
rain forest. Tropical, shady, wet. Alt. 300 m. Height of plant 5-6
m. Flowers cauline. Rose-white interior ; red appendages 10 cm. long;
corolla deep red. Grows deep in rain forest. Leaf and petiole with
hairy brown wool.’’ February 25, 1942, Univ. California 3rd Bot. Gard.
Exped. Andes 1942, 30173 (Leg. R. D. Metcalf & J. Cuatrecasas).
Ecuapor: “‘Foot of Western Cordillera. Alt. 100 m. In the shade
of the forest. Tree 5 m. high, 7 em. diameter. Bark blackish with
small warts. Leaves alternate, palmate, with 7 leaflets; leaflets 60 em.
long; leafstalks 60 em. long with 2 quite narrow, 24 em. long stip-
ules. Leaves and stem, when young, brownish velvet-hairy. Flowers
on the old stem, on short peduncles. Flower almost 3 cm. in diameter,
dark red, with 8 em. long reddish-white, thread-like appendages.
Fruit not seen, said to be cacao-like, with edible, white, sweet husks
of the seeds. Rare. Not used. Vulg. cacao de monte.’’ No date, A.
Rimbach 48.—Provincia Pichincha, Santo Domingo de los Colorados.
Alt. 800 m. ‘‘Especie arbores con flores directamente pegadas al
tronco. Flores rojas.”’ August 10, 1945, M. Acosta-Solis 10923.
Herrania purpurea (Pitt.) R. . Schultes in Caldasia
2, no. 9 (1944) 3383.
[ 132 ]
Theobroma purpureum Pittier in Fedde Rep. Sp. Nov.
13 (1914) 319.
The binomial Herrania purpurea was published as a
nomen nudum in the first edition of Thomas Belt’s **The
Naturalist in Nicaragua’’ (1874) 116. Belt wrote:
‘*About here grows a cacao (Herrania purpurea) differ-
ing from the cultivated species (Theobroma Cacao).”” I
have been unable to discover a description of the plant
or a publication of the binomial prior to 1874. In the
preface of his book, Belt stated that ‘‘Prof. D. Oliver
of Kew has kindly named for me some of the plants.”’
In the collection of Herrania at Kew, I did not find any
specimen from Nicaragua collected prior to 1874 and
annotated with this binomial. Dr. N. Y. Sandwith of
Kew has kindly searched through the archives and _ re-
ports that he can find nothing which might suggest that
Oliver had published the binomial.
Pittier’s description of Theobroma purpureum was
based upon a Panamanian plant collection, and he made
no mention of a prior publication of this specific epithet.
There is no doubt that the binomial which Belt published
refers to the same concept which Pittier later and inde-
pendently described and for which he used the identical
specific epithet. In accordance with the International
Rules of Botanical Nomenclature, therefore, we must
consider Pittier’s Theobroma purpureum as the first valid
use of the specific epithet.
'TERNSTROEMIACEAE
Mahurea tomentosa Ducke in Arqu. Inst. Biol.
Veg. Rio Janeiro 1 (19385) 208; in Trop. Woods 43
(1935) 22.
It would appear that this collection is the first of the
genus Mahurea from Colombia. There are several spe-
cies from Venezuela, the Guianas and Brazil, usually
[ 133 |
associated with the old Duida-Roraima type of flora.
Schultes 3929 was collected in a sandy caatinga at La
Chorrera, a locality having many elements in common
with the floras of the Apaporis, Vaupés and upper Rio
Negro caatingas.
CotomsiA: Comisaria del Amazonas, Rio Igaraparand, La Chorrera
and vicinity. “‘Bush. Flowers pink.’’ June 6, 1942, Richard Evans
Schultes 3929.
COMBRETACEAE
Combretum karijonorum P. 1. Schultes sp. nov.
Frutex scandens. Ramuli glabri et subnitidi, teretes,
2 mm. in diametro. Petioli robustiores, brunnei, 4 mm.
longi. Folia papyracea, atroviridia, ovata, plerumque
11-16 cm. longa, 8.5-11 cm. lata, basi rotundata, apice
breviter acuminata, integra, supra nitida, omnino glabra;
venis secundartis plusminusve septem supra haud impres-
sis, subtus prominenter elevatis, tertiis prominentioribus.
Inflorescentiae ex axibus superioribus prorumpentes,
usque ad 15 cm. longae. Flores adhuc ignoti. Samara
30 mm. longa, 12-183 mm. lata, omnino glabra, maturi-
tate rufescentia, alis elongato-linearibus, 2.5-8.5 mm.
altis, margine subintegra, transverse minutissime strio-
latis.
Combretum kariyonorum appears not to be closely al-
lied to other known Amazonian species of the genus and
‘an. be distinguished at once by its unusual fruit. The
samara is very long in relation to its lateral dimensions—
the body being elongate-lanceolate and the wings linear,
nearly as wide at the center as at the ends. This shape
is unusual in Combretum, There are other differences of
lesser importance, such as the extreme reduction of the
petioles which gives the large and ovate leaves almost a
subsessile appearance.
The specific epithet refers to the once populous and
fiercely warlike tribe of Karijona Indians, who formerly
[ 184 ]
inhabited the upper Apaporis basin, now reduced to
fewer than twenty individuals residing on the Rio Vaupés
and at La Pedrera on the Rio Caqueta. The Karijonas
refer to this plant as le-che-mah.
Cotomapia: Comisaria del Vaupés, Macaya-Ajaju River confluence.
Mount Chiribiquete. ‘‘Vine. Fruit green, with reddish tinge.’’ May
15-16, 1943, Richard Evans Schultes 5452.
Combretum rotundifolium L. C. Richard in Act.
Soc. Hist. Nat. Paris 1 (1792) 108.
In spite of its abundance in Amazonian Colombia,
Combretum rotundifolium does not seem to have been
reported from this Republic. Collections have also been
made in Venezuela and the Guianas.
CotomsiA: Comisaria del Vaupés, Macaya River, vicinity of Cachi-
vera del Diablo and mouth of river. Altitude about 300 meters.
‘*Flowers red. Vine.’? May 1943, Richard Evans Schultes 5501.
APOCYNACEAE
Ambelania cuneata Mueller-Argoviensis in Mar-
tius Fl]. Bras. 6, pt. 1 (1868) 17.
This represents apparently the second collection of
Ambelania cuneata. The type was from the upper Rio
Negro near the confluence of the Rios Guainia and
Casiquiare.
Cotompra: Comisaria del Vaupés, Rio Vaupés, Cafio Pact. Sandy
6é“ ; ’ =
or rocky savanna. Bush. Flowers yellowish.’’ March 6, 1944,
R. BE. Schultes 5815.
Ambelania zschokkeiformis Markgraf in No-
tizbl. Bot. Gart. Berlin 12 (1985) 295.
The two collections cited represent apparently the
second and third collections of this species, which was
described on the basis of material which Ducke found
on the Mauésmirim, an affluent of the Rio Maués. The
Marmellos material establishes the occurrence of A mbe-
lania zschokheiformis in the great Rio Madeira basin and
greatly amplifies the known range of the species.
[ 185 ]
Brazi.: Estado do Amazonas, Rio Marmellos (affluent of Rio Ma-
deira), Igarapé de Poté. ‘*Small treelet growing in dense stands on
sand bars. Trunk very stout at base, up to 18 inches in diameter
rapidly tapering upwards. Bark light grey-brown, corky. Flowers
cream. Latex white. Trunk always grows at 20° angle from vertical
against current. ‘Seen only in Marmellos’ according to inhabitants.
One of most characteristic sand bar plants. In rainy season almost
completely submerged. Common name: Oboia.’’ August 6, 1945,
Richard Evans Schultes & Edgar Cordeiro 6514.—Same locality, near
mouth of river. ‘‘Boia. On sand bars.’? August 12, 1945, Richard
Evans Schultes & Edgar Cordeiro 6533.
EXPLANATION OF THE ILLUSTRATION
Pirate XXVIII. PARAscHEELIA ANCHISTROPETALA
Dugand. Habit photographs.
Photographs by R. E. Scuutres
[ 186 ]
PLATE XXVII
EXPLANATION OF THE ILLUSTRATION
Prare XXVIII. (Upper figure). Herranta Koran-
orumM R. BE. Schultes. Habit photograph of flowers
of Schultes 3478,
Photograph by R. KE. Scuurres
(Lower figure). Herrania pULCHERRIMA Goudol var.
pacirica R, Eb. Schultes. Habit photograph of the
fruit.
Photograph by J. Cuarrecasas
PLATE N XVIII
EXPLANATION OF THE ILLUSTRATION
Prare XXIX. Herrania Camarcoana R.E.Schultes.
(Upper figure). Habit photograph of flowers of
Schultes & Pires 9130.
(Lower figure). Habit photograph of fruits of type
plant.
Photographs by R. Ei. Scuuvres
Say .
PIiATE ONS
EXPLANATION OF THE ILLUSTRATION
Prare XXX. (Upper figure). Herrania Mariar
(Mart.) Decaisne ex Goudot var. puruMAyonts R.E.
Schultes. Habit photograph of flowers and fruits of
type plant.
(Lower figure), AMBELANIA ZSCHOKKEIFORMIS Mark-
graf. A colony along the bank of the Rio Marmel-
los, Brazil.
Photographs by R. E. Schultes
PEATE X XN
EXPLANATION OF THE ILLUSTRATION
Prate XXXII. Raparea Mopestra Maguire. 1, en-
tire plant, about one third natural size. 2, spike-
let, somewhat more than one half natural size. 3,
bractlet of spikelet, somewhat more than one and
one half times natural size. +, calyx, semi-diagram-
matic, shown spread out in a single plane, some-
what more than one and one half times natural
size. 5, corolla, semi-diagrammatic, shown spread
out ina single plane, stamens attached in position,
somewhat more than one and one half times natu-
ral size. 6, stamen, front and side view, somewhat
more than twice natural size. 7, seed, adaxial face,
somewhat more than twice natural size.
PraATE A XX I
EXPLANATION OF THE ILLUSTRATION
Prare XXXII. Herranta Camarcoana R. FE.
Schultes. 1, leaf, about one quarter natural size. 2,
Hower, about one half natural size. 8, petal, about
twice natural size. 4, staminode and anthers, about
twice natural size. 5, ovary and style, about four
times natural size. 6, fruit, about one half natural
size.
Drawn by EB. W. Suivi
L ss
Disha
RE. Schuttes
EXPLANATION OF THE ILLUSTRATION
Prare XXXIII.
HerRANIA
Schultes.
KANUKUENSIS RR. Ff,
1, leaf, about one sixth natural size. 2,
fruit, about one half natural size. 8, portion of
under surface of leaflet, showing pilosity, about
four times natural size.
Drawn by EK. W. Siri
PLATE XXXIII
» HERRANIA Ranukuernsis
REC Bes
Lio
\
A
\
ON
EXPLANATION OF THE ILLUSTRATION
Prare XXXIV. Herranra koranorum FP, E.Schultes
1, leaf, one half natural size. 2, flower, one half
natural size. 8, petal, about twice natural size.
1, staminode and anthers, about twice natural size.
5, ovary and style, about four times natural size.
Drawn by BE. W. Siri
PLATE XXXIV
HERRANIA hofamorum
RE. Schuttes
A
(SFeigey 7 Jats
pals
a“
ond
See.
Aen
eee
Ste
Pree
. YA Yl Wid
WBS
ae
3%:
EXPLANATION OF THE ILLUSTRATION
Prare XXNXV. Herrania nycreropenpron FP. FE,
Schultes. 1, leaf, about one quarter natural size.
2, Hower, one half natural size. 3, staminode and
anthers, about twice natural size. 4, ovary and
style, about four times natural size. 5, fruit, about
one half natural size. 6, portion of under surface
of leaf, showing pilosity, about four times natural
size.
Drawn by KW. Suivi
PLATE XXXV
hy
7 he,
ms
pegs ac be eS eager SSS a thine
a
Pe
AY
Ar
By
HERRANIA myctero de
RE. Schultes
wndron —
EXPLANATION OF THE ILLUSTRATION
Prare XXXVI. Comprerum Karwonorum PR, FE,
Schultes. Habit drawing of fruiting braneh from
the type plant, natural size.
Drawn by Kh. W. Suivi
| es ee, |
Jy hike: <<
WN
/ Y
4 4 \
y} J ih
\\
\
\\K\\\
\h,
AK
CA
}
722
COMBRETUM
so -
Fe
APUIOMOPUML
\
ee
BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY
VoL. 14, No. 6
CamBripGr, Massacuusetts, NoveMBerR 6, 1950
NOTES ON PERUVIAN CINCHONAS—I
BY
W.H. Hover’
THE numerous collections of species of the genus Cin-
chona made throughout the Andes during the recent
war by botanists of the Foreign Economic Administra-
tion (formerly the Office of Economic Warfare) have
made it possible to reexamine critically the nomencla-
ture and taxonomy of this admittedly difficult group.
During the past century, dozens of species were described
in the genus, but Standley (in Field Mus. Bot. 6 (19386)
24-33) claims that many of these ‘‘pertain to forms of
scarcely or not at all more significance botanically than
horticultural varieties of common garden vegetables. ”’
Therefore, in the most recent treatment of the genus
for Peru, Standley (zbid.) reduced the number of Peru-
vian species to a mere handful, admitting, at the same
time, that additional material might make it possible to
amplify his treatment.
As cinchona botanist in Peru during the period 1943-—
1945, the writer had the unusual opportunity of collect-
ing and seeing in their native haunts practically all the
important forms of Cinchona occurring in that country.
As a result of this fieldwork and subsequent herbarium
' Associate Professor in Botany, University of Massachusetts (Am-
herst); formerly Botanist in charge of scientific surveys, Cinchona
Mission, United States Office of Economic Warfare, Lima, Peru.
[ 137 ]
study, it is his opinion that the number of species listed
by Standley for Peru, and probably elsewhere, may be
increased, for among the innumerable forms, varieties,
and hybrids represented in the enormous literature of
the genus, there are certain well-defined concepts which
appear to be more or less stable and are easily recognizable
especially in the field. Kven among such a notoriously
variable assemblage as is contained in the complex of
Cinchona pubescens Vahl sensu lat., there are entities
which appear to be very stable. As an example, one
might select the tree known as C. succirubra Pavon
ex Klotzsch which was originally collected by Richard
Spruce in Ecuador. This tree wherever grown, in the
Far East, in Guatemala, and elsewhere, although often
under ecological conditions quite different from its home-
land, remains apparently true to type, not only in its
general morphological characteristics, but also in such
physiological features as the patterns of alkaloidal yield.
This is understandable, for the Andean terrain, like
any lofty highly-dissected mountain mass, offers an ex-
traordinary range of very localized and disjunct habitats
which have permitted active speciation to occur. Cer-
tainly if adjacent Andean hoyas, isolated as they often
are from their neighbors, can be marked by considerable
local endemism among some groups of plants, why then
cannot they be so considered as to their cinchonas as
well?
The writer feels that certain entities at present con-
sidered as synonyms in the genus Cinchona were reduced
to synonymy often without sufficient study and clearly
must be resurrected if any semblance of order is to come
out of this chaotic group. Rusby (in Bull. Torrey Bot.
Club 58 (1981) 523-5380) already has protested Standley’s
recent treatment of the genus for Bolivia (in Field Mus.
Bot. 4 (1981) 266-278), and the present writer agrees
[ 138 ]
with him in part, at least regarding the status of certain
species such as Cinchona Calisaya Weddell which he has
met with in his travels in southern Peru. Indeed, the
writer cannot but feel that a great deal of careful taxo-
nomic work was accomplished by cinchona botanists of
the past centuries and especially by such men as Ruiz
& Pavonand Weddell who had become well acquainted
with the species in the field.
In the pages that follow, the writer initiates a critical
discussion of some poorly known Peruvian cinchonas in
the light of his observations in the field. The species
treated in this paper are easily recognizable because of
the rather dense covering of hairs found on the leaves
and young growth. Our knowledge of these species, up
to now, has been based for the most part on the most
meagre and fragmentary of herbarium material; in fact,
certain of them are known only from types collected
more than a century ago. Inasmuch as the existing de-
scriptions of these plants are either incomplete or based
only on the original collection, the writer includes modi-
fied descriptions based on a study of recent collections.
Besides his own collection (WHH), the first set of
which is deposited in the herbarium of the United States
National Arboretum (USN A), the writer has examined
specimens of cinchonas in the following institutions, to
whose curators he is indebted for kindnesses received
during the study: Chicago Museum (F), Gray Herbar-
ium (G), University of Massachusetts (M), New York
Botanical Garden (N Y), Museo Historia Natural ‘‘Javier
Prado”’ of Lima (L), the United States National Her-
barium (US), and the herbarium of the Estacién A gri-
cola de Tingo Maria in Pert (TM).
CINCHONA CARABAYENSIS Weddell
The identity of this cinchona has remained uncertain,
[139 ]
since it was incompletely described by Weddell (in Ann.
Sci. Nat. III. 10 (1848) 9) from fruiting material col-
lected by him in the Province of Carabaya (Dept. of
Puno). While collecting in the same area during 1948,
I discovered Cinchona carabayensis, but unfortunately
my specimens of this species were among a number lost
in transit in Peru. I still have very sketchy field notes
and these may be of some value in relocating the species.
In the ‘‘Flora of Peru’’ Standley states that C. cara-
bayensis was described ‘‘from thickets on the summits
of the mountains between the valleys of the Province
of Carabaya, especially near San Juan del Oro.’’ Pre-
sumably these data came from the type label, for Wed-
dell, in his original descriptions, merely gives the locality
as ‘‘Peruvia.’” San Juan del Oro, now long abandoned
and consequently not found on modern maps, was the
most famous of the Spanish placer mining centers in
this auriferous zone of southern Peru and was located
on the ridges separating the watersheds of the Upper
Inambari (here called the Huari-Huari) and Upper Tam-
bopata Rivers. The much overgrown site of the mine
may be reached by trail in about two days from the town
of Sandia. Since Weddell’s time the Province of Cara-
baya has been divided, with the result that old San Juan
del Oro, as well as most other important cinchona areas
(such as the Tambopata Valley, ‘‘Valle Grande’’ of the
Huari-Huari, ete.) inthe Department of Puno, are now
included in the modern Province of Sandia. On the
other hand, the modern Province of Carabaya, with Mac-
usani as the capital, occupies only that part of the old
Province of Carabaya lying west of a line drawn north
and south through Limbani, a line which roughly paral-
lels the Limbani-Mina Sto. Domingo-Astillero mule
trail.
It was in this same general area in early 1948 that C.
[ 140 ]
carabayensis was rediscovered. About a half day north-
east of Sandia the present trail into the eastern forested
country (called montana in Peru) forks into three trails,
one of which continues down the valley of the Sandia
River into the old San Juan del Oro area, another cuts
off over aridge to the east to enter the T'ambopata Val-
ley, and the third runs northwesterly over a very steep
ridge into the Valle Grande section of the Huari-Huari
Valley via the small Cachi-Cachi placer mine. The last-
named trail passes in its preliminary ascent over a very
steep ridge known as Ramospata which is covered with
thickets and small trees characteristic of the Andean
tree-line or ceja de la montana, In this thicket commu-
nity C. carabayensis, locally called echenique, is common
and is particularly abundant between the tambos (trail
shelters) known as Ichubamba and Ramospata.
Specimens of this cinchona are shrubs or occasionally
small trees seldom attaining a height of over three
meters. In its general habit the species resembles C.
Josephiana Weddell (also called echenique) which is
found at lower elevations on the same slopes of the San-
dia Valley. From glabrous C. Josephiana, C. carabayen-
sis may be separated easily by the presence of tomentum
which covers all young growth, twigs, inflorescence
branches, lower surfaces of the leaves, ete. From my
field notes is appended here a general, if still inadequate,
description of this tree transcribed in the field from tresh
specimens in the hand :—‘‘leaves averaging 15 cm. long
by 5.5 em. broad, similar in shape to those of C. Calisaya
Weddell but dull in aspect rather than lustrous and
smooth; petioles similar to those of calisaya; basal por-
tion of the midrib with a suggestion of red on the upper
surface; venation more complex than that of calisaya;
young leaves with scattered pubescence on upper sur-
faces, all leaves with tomentum beneath, the same type
[ 141 ]
of tomentum covering all parts of young growth includ-
ing inflorescences; flowers distinctly pink especially in
bud, fainter when open, the calyx a deeper pink chang-
ing into deep red as the young green capsule develops. ”’
In its densely tomentose habit C. carabayensis differs
from all other cinchonas known to me from southern
Peru. The closest relative of this rare tree of the Andes
of Carabaya appears to be the similarly hairy C. Pahu-
diana Howard (discussed below), a better-known tree
of the Chanchamayo Region of central Peru. These two
species have been considered conspecific by Standley, but
the writer’s impression from field observation is that they
are distinct. However, the fragmentary herbarium ma-
terial of C. carabayensis and Weddell’s rather incomplete
description make it impossible to compare critically these
two trees. Howard noted a similarity between the two
species, but pointed out correctly that one was normally
a shrub, the other always a tree; he also noted differ-
ences in the leaves. Until more abundant material of
C. carabayensis is in hand, it seems best to consider it
apart, especially since the two species are rather widely
disjunct. It should be pointed out that C. Pahudiana,
not C. carabayensis, was the species introduced into cul-
tivation in Java and elsewhere in the past century.
In addition to the living plants, the writer has also ex-
amined material, possibly the type (ex herb. Paris), in
the herbarium of the Chicago Museum (sheet no. 971880)
and collected by Weddell in the ** Province of Carabaya. ”’
Besides the differences noted above, the capsules of Wed-
dell’s specimen are much coarser and more woody than
those of C. Pahudiana. Seed differences may also exist,
for the ones examined are large in comparison with those
of most Peruvian cinchonas. It is hoped that botanists
who may get into this interesting area of southern Peru
will keep their eyes open for this neglected species.
[ 142 ]
CINCHONA GLANDULIFERA Ruiz & Pavon
As far as I can learn, this little-known species has not
been collected until recently, since it was originally
discovered and described by Ruiz and Pavon in central
Peru. Standley discusses Cinchona glandulifera in his
treatment of the genus in the Flora of Peru and states
that, besides the herbarium specimen at Berlin (probably
the type) which was destroyed during the war, material
(an isotype‘) is also to be found in the Delessert Herbar-
ium. Fortunately a photograph (Field Museum neg. no.
102) was made of the Berlin specimen and this agrees
very well with plates of this species published in the
works of Ruiz and Pavon and of Howard.
Recently my good friend, Ing. Hernan Augusto (a
Peruvian agronomist and Superintendent of Fundo
Sinchono, and formerly associated with the U.S. Gov-
ernment Cinchona Mission in Peru), made available ex-
cellent specimens and data of an unidentified cinchona
collected by him in the upper Monzon River valley near
Tingo Maria in 1944. His material proved to represent
Cinchona glandulifera, from the type locality, and was
an excellent match for the specimen of Ruiz and Pavon.
The type material of Ruiz and Pavon originated in the
mountains near Chicoplaya (‘‘in Peruviae Andium mon-
tibus nemorosis ad Chicoplaya runcationes, Carpales dic-
tas’). The species is also reported by them from Monzon,
Panatahuas, and Cochero (also written Cocheros or Cu-
chero).
The type locality of Chicoplaya is not shown on most
modern maps’; it is a hacienda in the Monzon valley
'Chicoplaya, type locality of Cinchona glandulifera, and nearby San
Antonio de Playa Grande, type locality of C. micrantha R. & P. ap-
parently were not visited by Ruiz and Pavon but by their collectors,
including Juan Tafalla, for the localities are not indicated on the map
of the itinerary of Ruiz and companions recently published (Field
[ 143 ]
about midway between Monzon and Tingo Maria. It is
the lowland terminus of a mule trail running down the
valley from the highland community of 'Tantamayo.
| have also collected this species in northern Peru near
the hamlet of Pomacochas in the Department of Ama-
zonas, and specimens referable to C. glandulifera were
collected in the same department by Antonio Raimondi,
and in the Department of San Martin by Richard Spruce.
The specimens from Amazonas differ somewhat from
typical and topotypical material in length of petiole and
in leaf shape, but otherwise are quite like the tree as it
occurs in central Peru. Inasmuch as C. glandulifera has
been found as far north in Peru as northern Amazonas,
it very likely occurs in the intervening territory, particu-
larly on the western slopes of those Andean ranges lying
west of the Huallaga River. This particular inaccessible
sector is actually one of the least known botanically in
Peru, at least as far as the genus Cinchona is concerned.
Unlike most other Peruvian species, C. glandulifera is
a shrub, in its type locality averaging three meters high
(Augusto!), and in the Pomocochas area seldom attain-
ing a height of over two meters. It is a sun-loving spe-
cies always occurring among other low-growing shrubs
and grasses in open communities known among the
Peruvians as pajonales. In Pomocochas this cinchona is
dominant in the community in which it occurs. Such
Mus. Bot. 21 (1940) 9). Ruiz and Pavon apparently got no farther
than Cuchero at the confluence of the Chinchao and Huallaga rivers.
The localities in question are both located in the Monzén River Val-
ley which joins the Huallaga at the site of present day Tingo Maria.
Both Chicoplaya and San Antonio de Playa Grande may be found
on the map (‘‘Plan del curso de los rios Huallaga y Ucayali y de la
Pampa del Sacramento’’) made by Manuel Sobreviela—one of several
maps in the atlas accompanying Herndon and Gibbon’s published
account of the exploration of the valley of the Amazon (Executive,
No. 53, 33d Congress, House of Representatives, Washington, 1854).
[ 144 ]
sites usually are to be found on open ridges generally
close to timber-line where tree types are replaced by
shrubby growth and herbaceous or suffruticose peren-
nials. Where pajonales merge into woodlands, types of
C. pubescens are found, and this species is of all the cin-
chonas the closest associate of C. glandulifera. In its
choice of sites and in its shrubby growth-form, hairy C.
glandulifera is identical with C. Josephiana Weddell of
southern Peru. However, the latter plant is easily dis-
tinguished from the former by well-marked morphologi-
cal differences, the most obvious being its glabrosity.
In central Peru this diminutive species with its nar-
rowly elongate panicles is in flower from at least Febru-
ary to July. Diagnostic features of C. glandulifera, other
than its shrub form and hairiness, are the very short cap-
sules and very small seeds.
An analysis of a composite sample of bark from several
individuals of this species from Pomocochas has yielded
only traces of erystallizable alkaloids, principally cincho-
nine. Even if the shrub were of value as a source of al-
kaloids, the small size of the stems would prohibit prof-
itable commercial exploitation.
There follows an expanded description of the species
based on recent collections.
Cinchona glandulifera Ruiz & Pavon FI. Peruv.
et Chil. 3 (1802) 1, pl. 224.
A shrub or oceasionally a small tree 2-5 m. high with
several trunks, these as much as 7 em. thick; bark ashy
to dark gray, the outer surface marked with inconspicu-
ous fine transverse fissures, the youngest branchlets pilose
or hirsute; stipules 12-40 mm. long, elliptical to oblong,
obtuse to acute, somewhat villose; leaves subsessile, or
with short petioles 2-15 mm. long, the blades ovate to
lanceolate or oblong-elliptic, 6-16 cm. long, 2.5-8.5 em.
[ 145 ]
EXPLANATION OF THE ILLUSTRATION
Prare XXXVII. Mule trail up the Ramospata ridge
out of the Sandia Valley (Dept. of Puno) Peru. In
the thickets in the foreground Cinchona carabayen-
sis Weddell was rediscovered.
Photograph by W. H. Hover
[ 146 ]
PLATE AXXVII
EXPLANATION OF THE ILLUSTRATION
Prare XXXVILTL. Cincnona GLANDULIFERA Rute &
Pavon growing near Pomocochas (Dept. of Ama-
zonas) Peru. Clump from which lodge 6/1 was
collected.
Photograph by W. HW. Hovae
PLhatE XXXVIILI
EXPLANATION OF THE ILLUSTRATION
Prare XXXITX. Cincnona Panupiana Howard. A
representative specimen (Hodge 6243) from the
region of the Chanchamayo Valley in central Peru.
Photograph by W. H. Hover
PLATE XXXIX
EXPLANATION OF THE LLLUSTRATION
Prare XL. Cinchona parasoricsa Pavon in thickets
near Tabaconas (Dept. of Cajamarca) Peru.
Photograph by W. H. Hover
PLATE XL
broad, the tips acute to obtuse, bases acute to obtuse or
occasionally subcordate, glabrous and sometimes some-
what lustrous above, hirsute beneath, especially on the
nerves, midribs usually reddish when fresh, vein-pairs
7-12; panicles generally narrowly elongate but small
and compact, leafy, hairy, terminal and axillary, many-
flowered, the flowers short-pedicellate, often in nodding
clusters; hypanthium about 1.5 mm. long, densely yel-
lowish-pilosulose; calyx darker, 1.5—2 mm. long with
short triangular teeth, pilosulose outside, glabrous
within; corolla pink, 7-12 mm. long, tomentose out-
side; capsule short, 7-15 mm. long, elliptical to oblong,
pubescent to glabrate; seeds small, about 4 mm. long.
COLLECTIONS EXAMINED:
Peru: Hudnuco: Described from the mountains of Chicoplaya,
Province of Hudmalies, Ruiz & Pavon; photograph of authentic ma-
terial (presumably the type) ex herb. Berlin (Field Museum negative
no. 102). Alturas de Carash (Province of Hudmalies), Monzén Valley,
on trail between Monz6n and Tantamayo, 1735 m., Augusto 8, 8A
(TM, WHH).—Amazonas: Summit of trail running between Pomo-
cochas and Yambrasbamba (Province of Béngara), 7200 feet, Hodge
6113 (L, M, USNA, WHH). Summit of ridge separating Pomocochas
from Shipasbamba (Province of Boéngara), 8500 feet, Hodge 6109
(L, M, USNA, WHH). Valle de Huayabamba (Province of Chacha-
poyas), 2500 m., Raimondi 974 (L, WHH fragment).—San Martin:
In monte Campana (Province of Lamas) prope Tarapoto (now San
Martin) Peruviae orientalis, R. Spruce 4832, Aug. 1856 (G, NY), cited
doubtfully as C. officinalis by Standley in the Flora of Peru.—Com-
mon names: cascarilla negrilla, cascarilla delgada, cascarilla del pajonal
(Huanuco), cascarilla negra (Amazonas).
CincHona PanupiaANa Howard
Although the status of C. carabayensis is uncertain
because of the lack of ample herbarium material, the
identity of C. Pahudiana is sure. This rather easily iden-
tified species was originally collected by Justus Charles
Hasskarl near Uchubamba which is a hamlet in the val-
ley of the Tulumayo River (near the Chanchamayo Val-
[ 147 ]
ley) in the Province of Jauja and Department of Junin.
Through the efforts of Hasskarl, living plants of C. Pa-
hudiana were established by the Dutch in their planta-
tions in Java, but, owing to the poor quality of the bark,
its cultivation was eventually abandoned. Howard has
given a full description and much historical information
about this species. The writer has collected ample ma-
terial of this tree which locally abounds in the moist
timberline forests of the upper Chanchamayo Valley
along the auto road between 'Tarma and San Ramon and
not far from the type locality. The description which
follows is based on these recent collections:
Cinchona Pahudiana Howard Ill. Nuev. Quinol.
Pavon (1862) pl. 21.
A small, often slender tree 5-7 meters high, 5-20 cm.
in diameter; bark thin, gray to brown’, the branchlets
pilose, the hairs usually shed below the second girdle ; sti-
pules 14-85 mm. long, 7-17 mm. wide, oblong-elliptical,
obtuse (rarely acutish), pilose on the outer surfaces;
leaves petiolate; the petioles 83-10 mm. long, pilose or
often densely tomentose; blades oblong-elliptical to
ovate or obovate, 8-28 cm. long, 4-15 cm. wide, the tip
rounded to obtuse, rarely acute, the base acute, decur-
rent onto the petiole, glabrate or sparingly hairy usually
on the veins above, generally yellowish-pilose below with
longer denser hairs on the veins, with 9-12 pairs of pri-
mary lateral veins; panicles many-flowered, terminal,
' Dried samples of bark have a strong lateral curvature and often
show narrow longitudinal ridges; there is also a strong tendency for
the outer cork layers to shed. Samples analyzed by the U.S. Govern-
ment Cinchona Mission Laboratory in Lima (as were all others men-
tioned in this paper) indicate that the species is of little value com-
mercially, a typical analysis of trunk bark of Hodge 6243 yielding
total alkaloids 4%, anhydrous quinine 0.53%, quinine sulfate 0.65%,
cinchonidine 1.02%, quinidine traces, total crystallizable alkaloids
28%.
[ 148 ]
the branches generally tomentose, leafy-bracteate at the
base, ultimate bracts linear, approximately 2 mm. or
less long; hypanthium 2 mm. long, densely tomentose ;
calyx 2 mm. long, sparingly pubescent to glabrate par-
ticularly at anthesis, dentate nearly to the middle, the
teeth erect, triangular-acute; corolla shell pink with a
deeper redder shade on outer surfaces, tube 12 mm. long,
limb 4 mm. long; capsule 10-80 mm. long, lanceolate,
becoming much broader in extreme dehiscence, pubes-
cent to glabrate, prominently 10-costate.
As stated above, C. Pahudiana is a small tree, never
shrubby like C. carabayensis. In general habit it more
closely resembles weedy forms of C. pubescens Vahl sensu
lat., and indeed it might perhaps better be included in a
subspecific category of that species. As with a number
of other cinchonas, the degree of pubescence among in-
dividual trees in a stand shows considerable variation and
this variability is especially to be noted in the juvenile
population. Nevertheless, C. Pahudiana in its typical form
seems to be easily identified and readily separable from
C. pubescens both in the field and in the herbarium,
because of its rather small, thick, generally tomentose
leaves with bases decurrent onto the short petioles.
COLLECTIONS EXAMINED:
This species is apparently of very limited local distribution in the
Department of Junin of central Peru from which area all specimens
have been collected. Photograph (Field Museum photograph no.
37218) of authentic (possibly type) Hasskarl material from Uchubamba
in the Province of Jauja. Province of Tarma, highway between Tarma
and La Merced, about 10-12 km. below Palca, 6000 ft., Fvinger 530
(L, USNA). Province of Tarma, low-statured mountain rainforests in
valley of Rio Palea at Utcuyacu midway on auto road between Palca
and San Ramon, 2000 meters, Hodge 6238, 6239, 6240, 6241, 6242,
6243, 6244, 6248 (L, M, USNA, WHH). Thickets and open woods,
Huacapistana (Province of Tarma), 1800-2400 meters, Killip § Smith
24214 (F, NY, US)... has unusually long capsules; cited by Stand-
ley in Flora of Peru as Ladenbergia Riveroana (Wedd.) Stand]. Pro-
[ 149 ]
vince of Jauja, Monobamba, 2000 meters, Raimondi 2959, 9326, 11180
(L, WHH (fragment) )'—Common name: cascarilla crespilla chica
(Hasskarl!, Raimondi!).
CINCHONA PARABOLICA Pavon ex Howard
In 1986, Standley described Cinchona Delessertiana,
basing his species on an unnumbered collection of An-
drew Matthews (in herb. Delessert) from Chachapoyas
(Department of Amazonas) in northern Peru. Comment-
ing on his new species Standley wrote: ‘‘It is hard to
understand why this Matthews collection, made long
ago, has not been named, but I can find no mention of
it in literature. The species is a well marked one charac-
terized by the very dense pubescence of short, spreading
hairs that cover all parts of the plant.’’ Although the
Matthews collection had remained unnamed since col-
lected, a binomial, C. parabolica Pavon, had been applied
to the same tree in 1862 when Howard published the
‘*Tllustrations of the Nueva Quinologia of Pavon.’’ So
Standley’s name must be reduced to synonymy.
Fortunately we have not only an ample description
(Pavon ex Howard) and a good illustration of the species,
but also what may be considered to be the type speci-
men of C. parabolica. 'The latter, located in the herbar-
ium at Madrid, was photographed by the Field Museum
(photograph no. 29688) and yields the following data
from the label: ‘*Cinchona parabolica, Cascarilla con hajas
rugosas, N. 562. L. 723. Ex Loxa.”’” Howard states that
his plate was drawn from this identical Madrid specimen.
A study of the ‘‘type specimen’’ at Madrid and of
Howard’s plate and the original description indicate
'These specimens of Raimondi, located in the Raimondi Herbarium
of the Museo de Historia Natural ‘‘Javier Prado’’ in Lima were ten-
tatively identified by me in Lima as C. glandulifera R. & P. (see Hodge,
W.H.: “‘Notas sobre los especimenes de cinchona del Herbario de
Raimondi’’ in Bol. Mus. Hist. Nat. (Lima) 9 (1945) 63-64).
[ 150 |
without doubt that C. Delessertiana and C. parabolica
are identical.
An earlier name, C. Mutisti var. 8. published by Lam-
bert (Ill. Cinch. (1821) 9) may be referable to C. para-
bolica, but this is impossible to confirm without an exam-
ination of the type. No mention is made by Lambert of
the collector of his specimen, but, like C. parabolica, it
was probably collected by Juan Tafalla. Lambert’s de-
scription is too brief for critical comparison with C. para-
bolica, but he does include in his diagnosis certain of the
important features of this species:
‘foliis. ... subtus ramulisque valde pilosis, margine un-
dulatis subrevolutis, panicula brachiata valde pilosa... .”’
and also under var. B. .. . folits ovalibus obtusis basi ro-
tundatis subcordatisve.”’
In 1849, Weddell took up Lambert’s species, C. Mu-
tisii, listing var. B. as var. crispa, but in 1869, Weddell,
combining this variety with C. parabolica, transferred
both entities to synonymy under another binomial of
Pavon’s, C. rugosa, a species from Cuenca, also published
by Howard. A summary of the tentative synonymy of
this cinchona and an expanded description based on re-
cent collections follow :
Cinchona parabolica Pavon ex Howard Il. Nueva
Quinol. Pavon (1862) pl. 16.
Cinchona rugosa Pavon ex Howard op. cit. pl. 17.
Cinchona Mutisii Lambert var. 8. [1]. Cinch. (1821) 9.
Cinchona Mutisu Lambert var. crispa Weddell in Ann.
Sci. Nat. LII, 11 (1849) 270.
Cinchona rugosa Pavon ex Howard var. crispa Wed-
dell in Ann. Sci. Nat. V, 11 (1869) 359.
Cinchona Delessertiana Standley in Field Mus. Bot.
6 (1936) 26.
A small tree, 6-8 m. high, up to 10 cm. in diameter;
[ 151 ]
the older fibrous bark light to dark gray or brown, marked
with short rather indistinct transverse fissures; branch-
lets thick, four-angled, with very dense brown pilose
hairs, internodes elongate; stipules erect, soon decidu-
ous, oblong, 12-35 mm. long with an obtuse tip, densely
hispidulose on the outer surface, glabrous on the inner
surface; leaves medium-sized, petiolate, subcoriaceous,
bullate or rugose, especially when fresh, margins con-
spicuously inrolled, particularly on the younger leaves;
petiole thick, 1-2 cm. long, densely hispidulose-tomen-
tose with brownish hairs ; blades oval-elliptical to oblong,
6-19 cm. long, 3-9.5 em. wide, with apex rounded or
obtuse and subapiculate, base obtuse to truncate, above
often at first densely appressed-pilose, later glabrate, with
veins and veinlets deeply impressed, below everywhere
densely hispidulose and with short loose brown to yel-
lowish hairs; midrib thick, usually reddish above for about
a third of its length, prominent below with approxi-
mately 12-16 pairs of conspicuous lateral veins; panicles
small to medium-sized (8-30 cm. across), terminal, at
least the younger branches densely hairy, leafy bracteate
at the base, many-flowered, the sessile flowers densely
aggregate, ultimate bracts linear, 3-4 mm. long; hypan-
thium 2-3 mm. long, densely fulvous-tomentose; calyx
3-4 mm. long, generally reddish, densely fulvous-tomen-
tose, toothed to the middle, the triangular teeth erect,
acute; corolla maroon (the color somewhat obscured on
the tube and outer surface of lobes by the dense ad-
pressed strigose hairs, but visible in the throat of the open
flower), 12 mm. long, with lobes much shorter (2 mm. )
than tube; capsule up to 25 mm. long, woody with a
rough-pubescent outer surface, seeds 7-8 mm. long, nar-
rowly elliptical.
I made several collections of C. parabolica in northern
Peru during the period 1948-1945, and some of my field
[ 152 |
observations may be pertinent. The species was first ob-
served growing just east of Huancabamba at Tabaconas
(in the Department of Cajamarca) which is some 200
kilometers (airline) northwest of the Province of Chacha-
poyas, the type region for C. Delessertiana. Subsequently
trees were also observed near Bambamarca (Cajamarca).
Although cinchonas were sought in the Chachapoyas
region, the species was never seen there. Mathews’ speci-
men undoubtedly originated not in Chachapoyas itself,
whose environs have long been depleted of forest, but
somewhere in the outlying fringes of that province. The
range of the species extends into the Loja region of
southern Ecuador, the data of Pavon published by How-
ard giving as localities ‘‘72 via collis Laterna, in collibus
Vilcabamba, Valladolid, locis altis frigidis Sabaneta, Que-
brada Onda et Cruz Grande nominatis, Provincia Laja.”’
Thus C. parabolica is known from the Departments of
Cajamarca and Amazonas (and undoubtedly eastern Pi-
ura) in northern Peru and extends northwards into the
Provinces of Loja and Azuay in southern Ecuador.
C. parabolica is a rather small tree, the largest example
seen by me reaching a height of about twenty-five feet
and a diameter of eight inches. The species is perhaps
the easiest of all cinchonas to recognize, because of the
very dense pubescence (noted by Standley and others)
which covers all parts of the young growth of the plant.
But even more characteristic in fresh material (though
often lost in pressed specimens) is the strongly bullate
or rugose nature of the subcoriaceous leaves whose mar-
gins often inroll or curl under conspicuously. ‘These
characters possibly account for the Spanish name crespilla
which is not only used today among cascarilleros of
northern Peru, but also is cited by Pavon in his descrip-
tions of C. parabolica and C. rugosa from Ecuador. The
thin, light-to-dark gray bark of this tree is quite fibrous
( 1538 ]
and the outer bark tends to show a very slight cross-
fissuring. This is usually a sign among cinchonas of a
valuable bark, but in this case it is almost completely
lacking in alkaloids (traces only), recent wartime analyses
made by the laboratory of the U.S. Government Cin-
chona Mission in Lima checking with the results ob-
tained by Howard in the last century.
In northern Peru, the moist highland forests where
cinchonas abound (6000-8000 feet), the so-called ‘‘cove
forests,’’ are isolated from one another by pajonales
(grasslands or cultivated areas) and are usually limited
to local guebradas. The ridge-margins of such forests are
made up of smaller second-growth species which merge
gradually and deeper in the quebrada with the taller
dominant species of the forest climax. C. parabolica is
a tree of the second-growth areas bordering these forests,
where it is often found intermixed with a weedy variety
of C. pubescens, called ‘‘amarilla,”’ and at higher eleva-
tions sometimes with the variety of C. officinalis known
as /qja. Within the climax forest, often only a quarter-
mile distant, large trees of colorada, another variety of
C. pubescens, may grow. Thus C. parabolica is in close
association, in northern Peru at least, with three other
cinchonas. Despite this, I have not observed any signs
of hybridization between C. parabolica and associated
species of the genus.
COLLECTIONS EXAMINED:
Ecuapor: Type of C. parabolica “*‘ex Loxa’’ (localities cited by
Howard include Laterna, Vileabamba, Valladolid, Sabaneta, Quebrada
Onda, and Cruz Grande); photograph of type ex herb. Madrid (Field
Museum negative no. 29638). Type of C. rugosa from the Province of
Cuenca (Pavon ex Howard). Type of C. Mutisii var. B. from the Pro-
vince of Loja (Pavon ex Lambert).
Peru: Amazonas: type of C. Delessertiana from the Province of
Chachapoyas, Mathews (herb. Delessert, Standley !). A specimen rep-
resenting C. Delessertiana, in the herbarium of the New York Botan-
[ 154 ]
ical Garden, bearing the label ‘‘Mathews, Peruvia, 1862’? probably
represents an isotype of Standley’s species. The date appears to be
incorrect, for Mathews died in 1841.—Cajamarca: Province of Jaen,
border of cove forest in ““Quebrada Pajonal,’’ Tabaconas, 7000-8000
feet, Hodge 6040, 6054 (L, M, USNA, WHH). Frontier trail be-
tween San Ignacio and Huancabamba, approximately 8000 feet, Hodge
6052 (L, M, USNA, WHH).—Common names: cascarilla crespilla,
(Cajamarca, Loja); Cascarilla crespilla con hajas rugosas (Loja).
ae
a
BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY
VoL. 14, No. 7
CaMBRIDGE, Massacuusetts, DecemBer 11, 1950
? ,
PREHISTORIC PLANT REMAINS
FROM BAT CAVE
BY
C. E. Smira, Jr.
Durine the summer of 1948, an archeological expedi-
tion under the direction of Mr. Herbert W. Dick of the
Peabody Museum of Harvard University discovered an
abundance of plant remains in a series of dry caves in
New Mexico. The maize remains in this material have
already been reported on (Mangelsdorf and Smith 1949).
The balance of the plant material will be discussed in
the present paper.
DESCRIPTION OF THE SITE
The site is known as Bat Cave. It is situated on the
edge of the Plains of San Augustin in Catron County in
west central New Mexico. The topography of the re-
gion has largely been determined by ancient Lake San
Augustin, which formed the caves. The lake basin is a
level expanse of fine water-laid material with occasional
spots of alkali in the playa. It is almost entirely sur-
rounded by mountainous terrain and is approximately
60 miles long and 6 to 20 miles wide.
The Bat Cave site consists of a large rock shelter (75
feet by 100 feet) with a series of smaller caves at one
side. ‘The smaller caves are 10 to 80 feet deep, and about
10 to 15 feet from the present floor to the roof near their
[ 157 |
mouths. All of the prehistoric material described below
came from these small caves.
PRESENT VEGETATION
A description of the present vegetation of the area
will serve as a basis for comparison with the material to
be described from the deposit. Near Bat Cave, the plains
consist of a large playa supporting a sparse growth of
four-winged salt bush (Atriplex canescens (Pursh) Nutt. ),
with areas of Heliotropium curassavicum LL. and Sporo-
bolus airoides (‘Torr.) Torr. The transition zone from the
plain to the fossil lake terraces is marked by Russian
thistle (Salsola pestifer A. Nels.). On the terraces Boute-
loua hirsuta Lag. forms a thicker ground cover than is
present on the plains. Numerous plants of the groundsel
(Senecio longilobus Benth.), stickseed (Lappula Redow-
sku (Hornem.) Greene), stickleaf (Mentzelia pumila
(Nutt.) Torr. & Gray), paintbrush (Castilleja minor A.
Gray), mock pennyroyal (Zedeoma Drummondi Benth. )
and verbena (Verbena ciliata Benth.) show that this area
is relatively free of alkali. In the rocky areas of the ter-
races a shrubby vegetation is present. M’allugia paradoxa
(D. Don) Endl. (Apache plume) is very common, while
Rhus trilobata Nutt. ex Torr. & Gray, Forstiera, Ly-
cium and Ribes spp. comprise the rest of the shrub flora
near Bat Cave.
The arroyos and canyons nearby are better watered
(although none have permanent streams) and conse-
quently the herbaceous vegetation is more varied. Nu-
merous members of the Compositae, Boraginaceae, Lab-
tatae, Leguminosae and other families were collected.
In no case, however, was there any extensive ground
cover formed by these plants.
At an elevation of 200 to 400 feet above the level of
the plains a distinct change in the vegetation is seen.
[ 158 ]
Pinus edulis Engelm. and Juniperus spp. are dominant,
while Bouteloua hirsuta Lag., Bouteloua curtipendula
(Michx.) Torr. and other grasses form an almost contin-
uous ground cover with scattered herbaceous plants and
‘acti (mainly Opuntia spp.). In one canyon near Bat
Cave there are several trees of Pinus ponderosa Doug}.
ex P. Laws. No other occurrence of ponderosa pine was
seen near the site.
During the dry season the vegetation has a distinctly
xerophytic aspect emphasized by the scattered plants of
Opuntia and Mammillaria. Only the heliotrope on the
plains and the shrubs on the terraces remain green. The
dark green of the pifon and juniper contrasts with the
brown ground cover.
With the advent of the summer rains in July and
August, many short-lived herbs appear over the plain
and hills making the entire area green. Most of the
vegetation blooms during this period. Heavy rains in
the hills and mountains usually fill the canyon beds with
temporary streams for several hours each day, and stand-
ing pools form on the playa.
EXCAVATION
The vegetal material recovered from Bat Cave was
collected during the course of excavation for archeologi-
cal remains. Measured areas were carefully dug by hand
troweling in twelve inch levels from the top of the de-
posit. The material of botanical interest was removed
and packaged. These arbitrary strata have no relation to
any archeological strata which may exist in the Bat
Cave deposit. Many cubic yards of material were re-
moved so that a fair sampling of the entire deposit was
obtained. Vegetal artifacts, to be described later by Mr.
Dick, were packaged separately. Aside from the maize
remains previously studied, no attempt was made in the
[ 159 |
field to separate cultivated plants from wild species, al-
though the specimens of beans and squash recovered
were obviously cultivated.
DETERMINATION OF THE REMAINS
All of the determinations are based upon characters
of taxonomic significance. Fragmentary material has
often been given only a generic name due to lack of the
detail needed for specific determination. A small amount
of the material proved indeterminable.
The identification of the wood and charcoal remains
presented certain technical difficulties in handling the
material. An unsuccessful attempt was made to soften
the charcoal enough to permit sectioning. In most cases
there was complete disintegration in the bleach solution.
Some few samples did not crumble, but these did not
soften sufficiently to be sectioned.
The wood remnants were softened by boiling. Because
of partial disintegration and chemical change, it was
found that no further softening by hydrofluoric acid
treatment was necessary. Although some material was
left unstained, the bulk of the wood sections were
stained by Heidenhain’s haematoxylin and safranin be-
fore being permanently mounted.
Except for the wood of the conifers and Quercus, the
only tree genera represented, the material could only be
identified after comparison with a set of slides made of
known woods occurring at similar elevations today.
Much of the old material represents such families as the
Chenopodiaceae, Rosaceae, Rutaceae, Rhamnaceae, and
Compositae.
The author wishes to express his sincere thanks for
the aid and advice accorded him by Professor Paul C.
Mangelsdorf, Professor Ivan M. Johnston, Professor
EKlso S. Barghoorn and the many others who have as-
[ 160 |
sisted in the identification of the material. Especial
thanks are due to Dr. S. F. Blake who identified the
Compositae. Without this expert help, much of the vege-
tal material would have proven indeterminable.
CULTIVATED PLANTS
There is no evidence that the Bat Cave people culti-
vated any of the native vegetation, although species such
as Yucca may well have been planted near the cave to
ensure a regular supply of fiber and amole. Many of the
herbaceous plants could have been grown with some suc-
cess, but the bulk of the material was probably prevalent
enough in the neighboring area to be easily collected as
the need for it arose.
Cucurbita Pepo Linnaeus Sp. Pl. (1753) 1010.
Aside from the maize, the only obviously cultivated
plants (that is, plants which are foreign to the region,
and which show increased size and other marks of culti-
vation) are squash and beans. All of the cultivated squash
remains are referable to Cucurbita Pepo L. The material
was found in every level (1 to VI) and comprises seeds,
rind and stems.
The seeds of Cucurbita Pepo are all characterized by
the moulding near the edge of the seed. Identification
is possible because of the difference in texture between
the center and the margin of all of the specimens. In
specimens without discoloration, there is no difference
in color in these areas. The seeds apparently all represent
the same variety of C. Pepo, as there is no distinct dif-
ference apparent in any of them. Furthermore, there is
no apparent size variation, as was shown by the maize,
from the bottom to the top of the deposit.
[ 161 |
Squash seeds
Level Number seeds Aver. width Aver. length
I 2 9.5 mm. 16.0 mm,
II 3 9.9 mm. 15.6 mm.
III 4 9,0 mm. 15.0 mm.
IV 11 9.0 mm, 15.8 mm,
V 8 8.9 mm. 14.9 mm,
VI 9 9.1 mm. 16.4 mm.
Tora Aver, 82 9.2 mm. 15.6 mm.
The average seed length is 15.6 mm., the largest strata
average being 16.4. mm. for stratum VI. The average
seed width is 9.2 mm., the largest strata average being
9.9 for stratum II.
Many pieces of rind were recovered from the excava-
tion, none of which were sufficiently large to permit an
accurate reconstruction of the size of the fruit. These
remains show no significant differences in thickness or
appearance. ‘The surface texture is generally smooth, a
few pieces showing slightly raised areas. There is promi-
nent veining on the inside of all of the specimens. The
thickness varies from about 1.5 mm. to about 3.5 mm.,
the average thickness being about 2.5 mm. Several pieces
of rind showing the flower attachment were found. These
were always concave in form, indicating that the fruit
was probably the shape of our modern pumpkin.
Three stems were found in the course of the excava-
tion. These are typical Cucurbita Pepo stems, flaring at
the point of attachment with the fruit, pentagonal in
shape and regularly fluted. Some idea of the size of the
fruits may be gained from the size of the stems.
Squash stems
Level Number stems Length Diameter Diameter
(upper end) (lower end)
Il 1 4.3 cm. 1.8 em. 3.0 cm.
IV 1 7.6 com, 2.0 em, 3.3.cm.
Vv 1 6.0 em. 1.8 cm. 2.8 cm.
[ 162 ]
The oldest stem was found in stratum [I and was ap-
parently cut from the fruit while still green. The length
of this specimen is 4.8 cm., with a diameter at the upper
end of about 1.8 em., and a diameter at the lower end
about 83cm. A whole stem, recovered from stratum LV,
is 7.5 em. long, 2 cm. in diameter at the upper end and
3.3 cm. in diameter at the lower end. The third speci-
men from stratum V is about 6 cm. long with a diameter
of approximately 1.8 cm. at the upper end, and 2.8 cm.
at the lower end.
Phaseolus vulgaris Linnaeus Sp. Pl. (17538) 728.
Only a few specimens of the common red bean were
recovered. ‘These have a smooth red coat with a white
hilum or eye. There is no marked difference in size or
appearance from level to level.
Common beans
Level Number beans Average width Average length
III q 8.5 mm. 13.0 mm,
IV 4 7.1 mm. 11.1 mm.
VI 1 8.0 mm. 11.0 mm.
The largest beans and the earliest in age were the two
specimens from stratum III (average width 8.5 mm. and
average length 18 mm.). In stratum IV the beans aver-
aged 7.1 mm. in width and 11.1 mm. in length. From
stratum VI a single bean 8 mm. wide and 11 mm. long
was recovered. A buckskin bag of beans was found in
stratum V, but these beans were not available for study.
The apparent decrease in size upward in the deposit is
probably due largely to the lack of sufficient material
for comparison. There is no ready explanation for the
lack of beans in strata I and II where both maize and
squash are well represented.
Contrary to the evolutionary series presented by the
Bat Cave maize, neither the cucurbits nor the beans show
[ 1638 ]
any marked developmental trend. The material available
for study does not furnish the detail found in the maize
remains, and there is little variation discernible in the
squash seeds and beans. ‘The number of squash stems is
also insufficient to furnish much information, Further-
more, the squash rind is devoid of any characteristic de-
tail. From the material studied it may be concluded that
at the time maize was first cultivated in a primitive form
at Bat Cave, C. Pepo was cultivated for its rather large
fruit. This squash apparently changed little if at all, while
the maize underwent a distinct development. Phaseolus
vulgaris does not appear in the oldest third of the Bat
Cave deposit. There is no apparent change in character
in the beans. It may perhaps be significant that the first
beans appear in stratum III in which the first teosinte
introgression is discernible in the maize.
Helianthus annuus Linnaeus Sp. Pl. (1753) 904.
Throughout the entire deposit flowering heads of
Aclhianthus annuus L. occur. There is no indication that
sunflowers were actually cultivated for their seed, al-
though they may well have been, as in other parts of the
southwest. The heads are of a size similar to those of the
wild plants which are very common in better watered
parts of the region today. These heads are about 2.5 to
3.5 cm. in diameter, and they vary in no way from the
bottom to the top of the deposit.
UNCULTIVATED PLANTS
The remainder of the Bat Cave vegetal remains may
be assumed to be uncultivated, as they are mainly ma-
terials which could easily have been gathered in a wild
state in the vicinity of the Cave. Descriptions are listed
in taxonomic sequence according to the Engler and
Prantl system, rather than according to abundance or
[ 164 |
supposed importance in the deposit. It is felt that this
will be a more usable plan for reference.
PINACEAE
Pinus edulis Hngelmann in Wislizenus Mem. North.
Mex. (1848) 88.
Pinon nuts were one of the most abundant plant re-
mains found in the Bat Cave deposit. Many hundreds
of broken shells were unearthed with a few whole nuts,
several cones and an occasional twig.
In strata I and II this species is represented by nuts
and cone scales. The remains in stratum III include a
twig with needles attached, as well as cones, cone scales
and seeds. From stratum LV pinion is represented by
both seeds and cone scales. In both strata V and VI
abundant seeds, cones, cone scales and twigs were re-
covered.
Thus, P. edulis is represented during the entire period
of occupation of Bat Cave.
Pinus ponderosa Douglas ex P. Lawson, Agr.
Man. (1836) 854.
Remains of this tree were found only in strata V and
VI. In stratum V ponderosa pine is represented by the
ends of two twigs with the needles adherent. A number
of separate needles and a cone were recovered from stra-
tum VI.
Pinus sp.
Wood of a species of Pinus was found in all strata
except I.
Juniperus pachyphloea Torrey in U.S. Rept.
Expl. Miss. Pacif. 4 (1857) 142.
Remains of juniper first occur in stratum II where the
[ 165 ]
plant is represented by large seeds. These may not be
seeds of J. pachyphloea, although the resemblance is close.
There were no remains identifiable as juniper in stratum
ILI. The seeds and berry found in stratum IV are char-
acteristic of J. pachyphloea, as are the numerous seeds,
berries and twigs bearing leaves found in both strata V
and VI.
Juniperus sp.
This genus is represented by wood fragments from all
but the first two strata.
The large amount of juniper remains in the upper three
strata contrasts sharply with the few seeds from stratum
Il and the lack of material from strata I and III. Using
pifon remains as a criterion, there must have been juni-
per trees in the area for the entire time of occupation,
so their absence in the lower parts of the deposit should
be attributed to disuse rather than to lack of available
material.
‘Ty PHACEAE
Typha latifolia Linnaeus Sp. Pl. (17538) 971.
Remains of aquatic or marsh plants are relatively rare
in the Bat Cave deposit. A portion of cattail leaf was
found in stratum LV. The largest amount of cattail ma-
terial, however, came from stratum V where numerous
pistillate heads and a small staminate head of 7' latifolia
were recovered.
At the present time, cattails grow only in recently
made ponds at Jacks Spring and 'T.U.T. Spring twelve
miles across the plain from Bat Cave. The abundance
of the cattail remains therefore suggests that during the
period represented by strata IV and V there was a per-
manent body of shallow water nearby. Probably the
large playa in front of the cave was at that time supplied
[ 166 }
with sufficient summer rain and winter snowfall to sup-
port a shallow water vegetation.
GRAMINEAE
Festuca Kingii (S. Wats.) Cassidy in Colo. Agr.
Expt. Sta. Bull. 12 (1890) 36.
Fruiting heads of this grass were found in stratum VI.
Poa Fendleriana (Steud.) Vasey in U.S. Dept.
Agr., Div. Bot. Bull. 18, pt. 2 (1898) pl. 74.
This grass was found in levels IV, V and VI of the
Bat Cave deposit.
Poa secunda Pres! Rel. Haenk. 1 (1830) 271.
A sample of P. seeunda was among the grasses re-
covered from strata III and LV.
Elymus salina M. FE. Jones in Proc. Calif. Acad.
Sci. ser. 2, 5 (1895) 725.
Fruiting material was found in stratum VI.
Sitanion Hystrix (Nutt.) J.G. Smith in U.S. Dept.
Agr., Div. Agrost. Bull. 18 (1899) 15.
This grass was also found represented only in stratum
VI.
Trisetum Wolfii Vasey in U.S. Dept. Agr. Month-
ly Rept. Feb., March (1874) 156.
Several specimens of 7. Wolfii were found in strata
Vaan V1.
Calamagrostis inexpansa 4. Gray Gram. and
Cyp. 1 (1834) no. 20.
C. inexpansa was found only in strata V and VI.
[ 167 ]
EXPLANATION OF THE ILLUSTRATION
Prare XLI. (Upper fig.). Plains of San Augustin,
New Mexico. Numerous clumps of saltbush (4tri-
plex canescens) thinly interspersed with grasses and
a few herbaceous plants. In the middle foreground
is the large playa in front of Bat Cave. Elevation
about 6500 feet.
(Lower fig.). Juniper (left) and pifion on the hills
directly above Bat Cave at an elevation of about
7000 feet. Ground cover here is heavier with
grasses, herbaceous plants, and cacti.
PLATE XLI
ne ee ee
EXPLANATION OF THE ILLUSTRATION
Prare XLIT. Remains of cultivated squash (Cucur-
bita Pepo) and beans (Phaseolus vulgaris) from the
deposit. A, stems from strata I], [Vand V. B,
specimens of rind from various strata. Note the
concave basal portions. C, squash seeds and beans
from strata as marked. D, fragment of rind of the
native wild cucurbit (Cucurbita foelidissima) with
attached seeds.
Pratre XLII
EXPLANATION OF THE ILLUSTRATION
Prare XLII. Gymnospermae and Monocotyle-
doneae from the Bat Cave deposit. A, Pinus. B,
Juniperus. C, Typha latifolia. D, Scirpus. K, Gram-
ineae. Fy Yucca.
PLateE XLIII
EXPLANATION OF THE TLLUSTRATION
Prare XLIV. Dicotyledoneae from the Bat Cave
deposit. A, Juglans major. B, Quercus. C, Amaran-
thus. I, Suaeda. K, Opuntia. F, Helianthus annuus.
PLATE XLIV
Muhlenbergia rigens ( Benth.) Hitchcock in Journ,
Wash. Acad. Sci. 28 (1983) 4538.
In strata LV and VI specimens of this grass were
found.
Sporobolus airoides ( 7orr.) Torrey in U.S. Rept.
lixpl. Miss. Pacif) 7 (1856) 21.
Abundant remains of this grass were found throughout
the deposit from stratum TE upward.
Sporobolus Wrightit Mianro cx Scribner in Bull.
‘Torrey Bot. Club 9 (1882) 103.
Specimens of stems were found from stratum EET up-
ward, Fruiting heads of SS. Wrightu were tound only in
stratum VI.
Bouteloua gracilis (J7BK.) Lagasca cv Steudel
in Nom. Bot. ed. 2, 1 (1840) 219.
The remains identified as B. eracilis were found in
strata TPE and TTT.
Bouteloua hirsuta Lagasca Var. Cienc. 2, pt. 4
(1805) 141.
This is one of the most common grass remains from
Bat Cave oceurring from stratum EE upward.
Sorghastrum nutans (l..) Nasi in Small, FL.
Southeast. U.S. (1908) 66,
One specimen of this grass was recovered from stra-
tum V.
Although much material representing the Grammeae
was found, there is little which may be interred from the
material at hand. The prevalence of Sporobolus airoides
and Bouteloua hirsuta throughout the deposit from stra-
tum TL upwards emphasizes the continuity in vegetation
[ 168 |
in the area for the last several thousand years. No typi-
cally aquatic grasses were found. None of the grasses
represented seem to have been gathered for food, so that
a continuous maize economy probably furnished the en-
tire grain supply for the Bat Cave people.
CYPERACEAE
Scirpus Olneyi 4. Gray in Boston Journ. Nat.
Hist. 5 (1845) 238.
A complete inflorescence as well as several detached
heads of §. Olneyi were found in stratum V.
Scirpus validus Vah/ Enum. PI. 2 (1806) 268.
A fruiting head of this bulrush was also found in stra-
tum V.
Scirpus sp.
Fragments of the thick leaves of an aquatic plant ap-
parently a Scirpus, were recovered from stratum IV of
Bat Cave.
The presence of these typically aquatic plants, as well
as Typha latifolia, in strata LV and V would seem to be
conclusive evidence for the presence of a permanent, al-
though perhaps shallow, body of water in the area of
Bat Cave during the period of occupation.
LILIACEAE
Yucca baccata Torrey in Emory Rept. U.S. &
Mex. Bound. Surv. (Bot.) (1859) 221.
Identifiable remains of this species were found in strata
LP sed tlt.
Yucca spp.
Numerous portions of yucca leaves, seed pods and
seeds were found throughout the deposit. In addition,
[ 169 |
two sections of rhizome, one from stratum IV and the
other from stratum VI, were recovered. Minor differ-
ences show that at least two species are present, but
lack of definitive characters makes impossible the accu-
rate identification of these remains.
SALICACEAE
Populus tremuloides Michaux FI. Bor. Amer. 2
(1803) 243.
Poplar leaves were recovered from stratum V. The
occurrence of this species with Typha and Seirpus fur-
ther emphasizes the probable occurrence of higher pre-
cipitation in the area of Bat Cave in former times.
JUGLANDACEAE
Juglans major (7Vorr.) Heller in Muhlenbergia 1
(1900) 50.
Numerous fragments of walnut shells occur through-
out the Bat Cave deposits.
FAGACEAE
Quercus Gambellii Nutta// in Journ. Acad. Nat.
Sci. Phila. ser. 2, 1 (1848) 179.
Leaf remains of Q. Gambellii were found in strata V
and VI.
Quercus grisea Liebmann in Overs. Danske Vidensk.
Selsk. Forh. 1854 (1854) 171.
This species is represented in the deposit by leaves
from stratum II upward.
Quercus sp.
Acorns were found in abundance in stratum III. A
single acorn shell was recovered from stratum V.
g
[ 170 ]
Wood of Quercus sp. was found to be most prevalent
in stratum III where six specimens were recovered. This
genus is represented by only one other wood specimen
from stratum VI.
It is not surprising to find Quercus remains in the Bat
Cave deposit, as oaks grow in the area today in the
better watered canyons. This further points to the great
similarity of the past vegetation of the area to that of the
present. It is unusual, though, that there should be so
many acorn shells and no juniper remains in stratum
III. At the time represented by this stratum there must
have been a much greater abundance of oak in the area,
which in turn would suggest a slightly higher rainfall
than that of today.
CHENOPODIACEAE
Chenopodium sp.
Remains belonging to this genus were found in stra-
tum IV.
Atriplex canescens (Pursh) Nuttall Gen. No.
Amer. PI. 1 (1818) 197.
Typical four-winged fruits of this plant were recovered
from strata IV and VI.
The shrubby members of the Chenopodiaceae are rep-
resented by the wood of Atriplex (canescens?) and other
woody material identifiable only to family. The former
is a very common shrub of the area today. The Atriplex
wood was found in stratum I, while the other material
came from strata II, ILI, [IV and VI.
Suaeda suffrutescens S. Watson in Proc. Amer.
Acad. 9 (1874) 88.
In strata I and V were found twigs of this plant.
[ 171 |
AMARANTHACEAE
Amaranthus blitoides S. Watson in Proc. Amer.
Acad. 12 (1877) 273.
One specimen was found in stratum V.
Amaranthus hybridus Linnaeus Sp. Pl. (1753)
990.
This species was found only in stratum I.
Amaranthus Powellii §. Watson in Proc. Amer.
Acad. 10 (1875) 347.
This is one of the most common plants in the deposit
and is represented by material from strata II, [11, V
and VI.
Amaranthus sp.
A specimen of Amaranthus from stratum IV was too
fragmentary for specific identification.
Plant remains representative of the centrosperms were
found repeatedly in the Bat Cave excavation, probably
because of their usefulness as food in an area where there
is no surplus of wild food plants. Here again similarity
to the present vegetation in the area of Bat Cave is
apparent.
SAXIFRAGACEAE
Ribes spp.
Ribes (aureum?) wood came from strata I, II and IV,
while two specimens of Rvbes, unidentified as to species,
were recovered from strata I and VI.
ROSACEAE
The family Rosaceae is represented by a single speci-
men of wood from stratum I.
[172 ]
LEGUMINOSAE
Oxytropis sp.
Seedpods of this plant were found in stratum V.
RUTACEAE
Ptelea sp.
A single specimen belonging to the genus Ptelea oc-
curred in stratum V.
RHAMNACEAE
Rhamnus sp.
Stratum V also yielded one specimen of the genus
Rhamnus.
CACTACEAE
Opuntia sp.
Numerous joints of prickly pear were found through-
out the deposit from stratum [II upward.
ASCLEPIADACEAE
Asclepias sp.
Milkweed stems and pods were found in quantity in
strata V and VI.
SOLANACEAE
Lycium sp.
Desert-thorn is represented by leaf remains from strata
IV, V and VI, and by twigs from stratum IV.
LABIATAE
Monarda sp.
Several fruiting heads of this plant came from stratum
"4 &
[ 173 ]
CUCURBITACEAE
Cucurbita foetidissima HBK. in Nov. Gen. et
Sp. 2 (1817) 123.
Remains of the fruit of this species were found in
strata IV and VI.
:
COMPOSITAE
Eupatorium sp.
Stems with leaves and involucral bracts were found in
strata LIV and VI.
Artemisia spp.
Numerous flowering stems of a sage brush were re-
covered from strata IV, V and VI. One specimen of
wood which is probably an Artemisia came from stratum
LY.
Pericome caudata 4. Gray Pl. Wright. 2 (1853)
82.
In stratum V a number of heads of this composite
were found.
Aplopappus gracilis (Nutt.) 4. Gray in Mem.
Amer. Acad. ser. 2, 4 (1849) 76.
In stratum VI two stems of this plant were recovered.
Verbesina sp.
Heads identified as belonging to this genus came from
stratum VI.
The composite family was well represented in strata
I, I11, IV and V by wood material identifiable only as
to family. ‘The composite remains are in no way unusual
as they represent only forms which grow in the area at
the present time.
Altogether the plant remains from Bat Cave comprise
[ 174 |
19 families and 89 genera. Although the selection of
these plants was entirely dependent upon the needs of
the people, it nevertheless provides us with a fairly sat-
isfactory record for several thousand years of the vege-
tation in the Bat Cave area.
As has been previously noted, the vegetation during
the period of the occupation of the Cave was remarkably
similar to the present day vegetation. Three species
present today are represented from stratum I to stratum
VI, and five additional species are represented from stra-
tum II to stratum VI. These plants include Pinus edu-
lis, Juniperus pachyphloea, Sporobolus airoides, Bouteloua
hirsuta, Juglans major, Quercus grisea, Amaranthus
Powellti and Helianthus annuus. All of these, with the
exception of the Juglans, are represented in the area
today. Moreover, species of Opuntia are represented from
stratum IIT to stratum VI, further emphasizing the sim-
ilarity to modern climatic conditions.
On the other hand, there are some evidences for a
slightly higher rainfall in the area during the Bat Cave
occupation. The persistent presence of Juglans major
may mean that it once grew in the vicinity. To the
author’s knowledge, the nearest walnut trees at the pres-
ent time are about 25 miles away in the Tularosa Moun-
tains. It could have been traded in just as easily. But
the presence of Typha latifolia in strata LV and V, and
Scirpus validus, Scirpus Olneyi and Populus tremuloides
in stratum V can only reasonably be explained by a
nearby body of water. At the present time, pools of
water remain on the playa for many days during the
summer rainy season. A better distribution of this rain-
fall throughout the year, together with a slight increase
in the average annual precipitation, would result in a
permanent shallow lake on the playa in front of Bat
Cave. The number of acorn shells in stratum ILI may
[175 |
also indicate a higher precipitation and thus a greater
abundance of oak in the area at the time of the stratum
LIL occupation. The necessity for an adequate water
supply during the period of Bat Cave occupation must
have meant a higher precipitation, as there is no perma-
nent source of water nearer than the springs at T.U.T.
Ranch and Jacks Peak about twelve miles across the
plain from Bat Cave.
At the present time flood water farming in nearby
canyons would be possible only in years in which the
rainy season started relatively early. In the summer of
1948, the rainy season did not begin until about July 15.
This probably would not permit a sufficient growing
period to mature maize, beans or squash before the fall
frosts at this altitude (6500 feet above sea level). Thus,
again it seems probable that the Bat Cave people had a
permanent water supply or that the annual precipitation
was more favorably distributed throughout the year.
The remainder of the Bat Cave plants are of no im-
portance as climatic indicators. All of the genera grow
in the area today, but their characteristics of growth give
no indication of past conditions.
From the nature of the plant remains we may assume
that the Bat Cave people practiced a subsistence agri-
culture supplemented by gathering wild plants as their
fruits matured, or as the need for various plants arose.
No evidence of extensive storage was encountered in the
excavation.
The subject of dating the Bat Cave deposit has been
purposely omitted thus far. Suffice it to say, Dr. Ernst
Antevs has studied the local geology. Although it was
reported in an earlier publication (Mangelsdorf and
Smith, 1949) that the cultural deposit from which the
plant remains were recovered could not have begun more
recently than 2500 B.C., Antevs now feels that this
[ 176 |
material could not have been deposited previous to about
2000 B.C. This point in time is established by the upper
limit of a thin dust layer upon which the plant-bearing
deposits rest. The upper limit has been tentatively dated
by Mr. Dick as 500 to 1000 A.D. These latter dates are
based upon pottery fragments from the upper 18 inches
of the Bat Cave deposit.
Additional evidence as to the age of this deposit is
also available. Vegetal remains from the Bat Cave de-
posit were sent by Professor P. C. Mangelsdorf to the
Institute for Nuclear Studies at the University of Chi-
cago where radio carbon dates were assigned. These vary
from 1752 + 250 for stratum VI to 2862 + 250 for stra-
tum II. The material from stratum I was insufficient to
obtain data. By extrapolation, however, it would appear
to be 8000 to 8500 years.
Thus, stratum II, for which the earliest date was ob-
tained, may contain material deposited as early as 1162
B.C. or as recently as 662 B.C. Should the earlier ma-
terial (stratum I) be 8500 years old, the deposit could
have been started about 1550 B.C. This date agrees
satisfactorily with the geological date furnished by Dr.
Ernst Antevs.
SUMMARY
1. Cultivated squash (Cucurbita Pepo) is present through-
out the Bat Cave deposit.
2. Cultivated beans (Phaseolus vulgaris) are found only
from stratum III upward, and then only in small
quantities.
3. Wild vegetal remains indicate semixerophytic vege-
tation in the past similar to the existing vegetation
near Bat Cave.
4. Remains of plants requiring a permanent body of
shallow water (Z'ypha latifolia, Scirpus Olneyi and
[177 ]
7
Scirpus validus) indicate a higher amount of precipi-
tation than at present for the time represented by
stratum V and possibly for stratum IV.
The presence of cultivated plants (maize, squash and
beans) and Juglans major also indicate a higher an-
nual precipitation during the time of the Bat Cave
occupation.
The abundance of acorns in stratum III may indicate
a higher precipitation at that time.
The Bat Cave people probably practiced a subsistence
agriculture supplemented by gathering wild plant
materials.
PLANTS OF BAT CAVE
—
—
_
—_
—_—
_—
—_
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=
_—
Zea Mays xX X
Cucurbita Pepo D4 xX
Phaseolus vulgaris
Helianthus annuus xX X
Pinus edulis xX xX
Pinus ponderosa
Pinus sp. (wood)
Juniperus pachyphloea
Juniperus sp. (wood)
Typha latifolia
Festuca Kingii
Poa Fendleriana
Poa secunda 5.4
Elymus salina
Sitanion Hystrix
Trisetum Wolfii xX X
Calamagrostis inexpansa xX
Muhlenbergia rigens
Sporobolus airoides xX
Sporobolus Wrightii
Bouteloua gracilis X
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II
Ill
IV
VA
Bouteloua hirsuta
Sorghastrum nutans
Scirpus Olneyi
Scirpus validus
Scirpus sp.
Yucca baccata
Yucca spp.
Populus tremuloides
Juglans major
Quercus Gambellii
Quercus grisea
Quercus spp. (acorns)
Quercus sp. (wood)
Chenopodium sp.
Atriplex canescens
Atriplex (canescens?) (wood)
Suaeda suffrutescens
Chenopodiaceae (wood)
Amaranthus blitoides
Amaranthus hybridus
Amaranthus Powellii
Amaranthus sp.
Ribes (aureum?) (wood)
Ribes sp. (wood)
Rosaceae (wood)
Oxytropis sp.
Ptelea sp. (wood)
Rhamnus sp. (wood)
Opuntia sp.
Asclepias sp.
Lycium sp.
Monarda sp.
Cucurbita foetidissima
Eupatorium sp.
Artemisia sp.
Artemisia sp. (wood)
Pericome caudata
Aplopappus gracilis
Verbesina sp.
Compositae (wood)
www
[ 179 ]
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LITERATURE CITED
Hitchcock, A. S. 1935. Manual of the grasses of the United States.
U.S. Department of Agriculture Misc. Publ. No. 200.
Kearney, T. H. and R.H. Peebles. 1942. Flowering plants and ferns
of Arizona. U.S. Dept. of Agriculture Misc. Publ. No. 423.
Mangelsdorf, Paul C. and C. E. Smith, Jr. 1949. New archeological
evidence on evolution in maize. Bot. Mus. Leafl. Harvard Univ.
13: 213-247.
Powers, William EK. 1939. Basin and shore features of the extinct
Lake San Augustin, New Mexico. Journ. Geomorphology 2: 345-
356.
Standley, Paul C. 1910. The type localities of plants first described
from New Mexico. Contrib. U.S. Nat. Herb. 13: 6.
Thompson, H.C. 1939. Vegetable crops. New York.
Whitaker, Thomas W. and G. W. Bohn (1950). The taxonomy, ge-
netics, production and uses of the cultivated species of Cucurbita.
Econ. Bot. 4: 52-81.
Wooton, E. O. and Paul C. Standley. 1913. Description of new
plants preliminary to a report upon the flora of New Mexico. Con-
trib. U.S. Nat. Herb. 16: 4.
Wooton, E. O. and Paul C. Standley. 1915. Flora of New Mexico.
Contrib. from the U.S, Nat. Herb. 19.
[ 180 |]
BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY
VoL. 14, No. 8
CamBripGeE, MaAssacuuseTTs, JANUARY 15, 1951
THE STORY OF PAN CHEWING IN INDIA
BY
M. Gowpa’
INTRODUCTION
‘There are so many ways of doing everything, all over
India.’’ This is particularly true of pan or betel chewing,
not only in India, but in all the Eastern countries where
pan has entwined the culture and conventions of the
people. Even the name pan has acquired a new conno-
tation in India. It originally meant betel leaf. In modern
writings and usage, however, pan refers to both betel
nut and betel leaf; thus in Hindustani pan dena means
the custom of giving betel nut and betel leaf, and in
South India tambula kodi conveys the same meaning.
The origin of its use fades into antiquity and is lost in
the dim past of the prehistoric periods.
The chief constituents of pan are betel nut, betel leaf
and quicklime, the inevitable three apices of the pan
triangle; Forster calls them ‘‘pan’s trinity.”” The other
materials used with them are merely ingredients.
The use of pan, a civility, a hospitality, a convention,
a habit, and an innocent after-meal breath-sweetening
practice, has often been treated in sweeping generaliza-
tions based on partial and imperfect facts. A practice
and habit that involves over one third of the human race,
‘Deputy Superintendent, Government Gardens, Lal-Bagh, Ban-
galore, India.
[ 181 |
in India, Burma, Ceylon, Malaya, Indonesia, the Philip-
pines, the Pacific Islands as far as Fiji and the Marque-
sas, China, Indo-China, Siam, East Africa, Arabia and
Persia, and is so intimately interwoven with millions of
individuals from birth to death, in joy and sorrow, in
respect and contempt, in friendship and enmity, and in
challenge and honour, must be looked into from all
angles.
Leaving aside the botanical and chemical analysis of
betel nut and betel leaf and the works of early writers,
the opinion expressed or written about pan by many re-
cent European writers has not been wholly true; the
custom has been exaggerated or represented as some-
thing abhorrent to Westerners. The reasons for these
sweeping generalizations, based on partial or imperfect
facts, that impressed me most on going through many
of these writings about pan are: (1) unfamiliarity with
the use of pan in any form; (2) judging with the pre-
conceived idea that anything the East does is something
strange, curious, mystic, or bound up with religion; (3)
lack of knowledge of the background of the culture of
the people and their food. Any one of these would be
sufficient to colour the author’s conclusions.
The reasons for the use of pan are lost in antiquity ;
one’s own ideas are as good as any others. In the East,
it is said to be because of the carminative effect, to keep
the gums and teeth strong, to aid digestion, to sweeten
the breath, or for something to chew after meals, or a
good practice by which to circumvent narcotic tempta-
tion. One of the very early writings, the Indian ‘‘Hito-
pades’’ (friendly advices), notes about thirteen properties
of pan; it removes bad odour; it expels phlegm; it ex-
pels flatulence ; it expels worms; it beautifies the mouth;
it helps digestion; it is sour; it is bitter; it is heating;
it is sweet; it is salt; it is astringent; and it excites de-
[ 182 ]
sire. Marco Polo speaks of it as “‘salutary.”*” One reason
that strikes me is this: coffee was introduced into India
only about 200 years ago, and tea might have been a
little earlier; but even now tea and coffee do not form a
part of all meals. Lunch or dinner or feast meals are ended
in many cases with curds, buttermilk, milk or sweets.
In all cases, however tasty or flavoured the food may be,
the food or beverage taste is gotten rid of from the
mouth after the mouth is washed. It is generally after
the meal and mouth-wash that pan or a piece of nut, or
a mixture of nuts is used to sweeten the breath. This
habit of the aftermeal use of pan or a piece of nut is the
most common practice that one comes across in India.
Garcia da Orta noticed this practice about 400 years ago.
‘*Among these people | Indians] it is so detested to smell
bad.”’ ‘‘Everyone chews it [pan] after meals.”’ ‘‘Many
Portuguese say that when they eat fish, they alternate
it with betel.’’ Forster writes, ‘‘their [pan chewer’s]|
breath is sweet; they are the exact antithesis of Italians,
and crowd for crowd I would rather be with them {pan
chewers ].”’
In all cases of pan chewing up to the beginning of the
17th century, and in the majority of cases now, the pan
was and is used to eat and swallow for its acquired taste,
and the colour in the mouth is rinsable, leaving no stain
on the teeth after washing.
After tobacco was introduced into India by the Por-
tuguese somewhere in the 16th century’ and its use in
' Baber in his Memoirs for 1519-1525, describing all the useful
plants and animals of India, makes no reference to tobacco. Garcia da
Orta lived in India from 1534 to 1564, and in his classical works on
Drugs and Simples of India, makes no mention of tobacco. “*Doubtless
to the Portuguese is due the credit of having conveyed both the plant
and the knowledge of its properties to India and China. It is said in
the Dara-shikohi that they had conveyed it [tobacco] to the Deccan
as early as 1508°’ (Watts). Asad Beg, in 1605, found some tobacco
[ 183 ]
India and China became widespread during the 17th
century, a new form of pan chewing developed, that is,
chewing betel nut and betel leaf with lime and tobacco,
but only to keep it in the mouth for its effect. Nothing
is swallowed or eaten in this type of chewing; the red
saliva is spit out as often as it collects in the mouth; the
mass of chewed pan with tobacco remains in the mouth;
finally, when the tobacco loses its effect, the whole mass
is spit out. This detestable habit of chewing has been
described as causing “‘the ghastly marks one commonly
sees in the streets.’’ The blackening of teeth is mostly
noticed among this class of pan chewers due to their
negligence in cleaning their teeth. In this method, the
pan is only an accessory or an accomplice to the tobacco
habit. All the low grade nuts and leaves are used up by
this class of chewers; the spicy and aromatic ingredients
used by the majority of chewers for taste are not used
by those who chew pan for the tobacco habit. Before
tobacco introduction, chewing was entirely to eat and
swallow, and spitting was rare, and even those who used
kath, spit out only the first saliva, if any; the rest was
chewed and eaten. Garcia da Orta has noticed this:
‘They spit out the first Juice... . some persons do not
do this but chew it all and afterwards take other leaves
in the same way.”’
Out of about 30 plant products which are used in some
form or other in chewing pan, only the betel nut (with
the exception of tobacco) contains alkaloids. here are
several of these, the most active principle being arecoline
C,H,,O2N (about 0.1%). Besides alkaloids, the nut
contains: Moisture: 81.8%; Proteins: 4.9%; Fats:
4.4% ; Carbohydrates: 47.2%; Minerals: 1.0%; Cal-
in Bijapur and took it to the Emperor Akbar who attempted to smoke.’’
.... Comes affirms that the seed cultivated in India in 1605 had been
brought from Brasil.’’ (Watts)
[ 184 ]
cium: 0.05% ; Phosphorus: 0.139% ; and a trace of [ron:
0.5 mg. per 100. Cured nuts are considered to be of the
highest quality. The curing improves the colour and taste
and removes the excess of tannin and mucilage in the
nut. Raw nuts contain 21.6 to 30.2% of tannin, whereas
cured nuts only have 8.6 to 15.1%. Cured nuts are more
commonly used in southern India than in northern India.
The betel palm and vine have been variously described
in Indian literature; the areca palm has been called ‘‘an
arrow from heaven’’; and poets have often symbolized
the palm and vine for enduring strength and beauty.
Hisrory, Borany AND CULTIVATION
a. Betel Palm: Areca Catechu L.
The genus name Areca is of South Indian origin.
Among the Nairs, Areca means ‘‘cavalier.”? In Kannada’
the betel palm is called the Adike tree, the nut Adike.
The specific epithet Catechu is probably from Kachu
meaning in Kannada ‘‘astringent substance. ’”’
The original home of the betel nut palm is uncertain, as
it is so widely cultivated in all the tropical parts of Asia
and in all the islands of the Indian and North and South
Pacific Oceans (see map). It has as many distinct names
as there are languages in the East. It is not found wild
anywhere in any of the countries of its cultivation. Its
numerous varieties, the many distinct names, and the
lack of wild individuals all indicate its antiquity, but do
not solve its origin. Several different opinions have been
expressed by botanists as to the original home of the
betel palm: Martius is uncertain, but thinks it is proba-
bly native to the Sunda Isles; Blume, Bretschneider and
Ridley consider it to be Malayan; Miquel and Blanco
‘Kannada is one of the major languages of South India. I am
not sure which is the older term, i.e., whether Areca is derived from
Adike or vice versa.
[185 ]
EXPLANATION OF THE ILLUSTRATION
Pirate XLV. A fruiting branch of the betel palm
(Areca Catechu L.) showing some of the fruits or
nuts in section. Betel nuts are one of the chief
ingredients of pan or betel chew.
From a painting by BLancur Ames.
[ 186 ]
PLATE XLV
do not seem to agree with them. Blatter claimed to have
received a letter from Fischer stating that he found in
the Attapadi Valley of Malabar, South India, the areca
palms growing wild in dense evergreen virgin forests.
Beccari’s reasoning seems to be more plausible; he con-
siders that Areca Catechu var. silvatica is very closely re-
lated to the cultivated palm, and this variety is found
wild only in the Philippines, and nowhere else. In addi-
tion to this variety, there are many Areca species found
wild in the Philippines which closely resemble the cul-
tivated plant. On these evidences, he thinks, the culti-
vated species probably originated in the Philippines.
The betel palm is a slender and graceful tree with a
strong columnar stem surmounted by a crown of pinnate
leaves. The straight, cylindrical, greyish trunk attains a
height of over sixty feet. From base to crown it tapers
imperceptibly and bears regular, faint rings of scars of
fallen leaves. The leaves are four to six feet long; the
lower portions of the petiole expand into a broad sheath
and cover the stem before falling off. The upper young
pinnae are confluent and stick out straight in the center
of the crown. The inflorescences are found on the stem
below the leaves, in the axes of old-fallen leaves; the in-
florescence is a spadix enclosed in a spathe and consists
of a much branched rachis bearing male and female flow-
ers. The fruit is ovoid or oblong according to the variety,
about the size of an egg, two inches thick and two and a
half inches long. Before ripening, the fruits are lustrous
green; when quite ripe, they turn to orange-yellow.
The pericarp is fibrous and husky. The seed, or kernel,
consisting of hemicellulose endosperm, is the ‘‘nut’’ used
for mastication. The tiny embryo at one end of the nut
drops off on drying or during the process of cooking.
The uncured (uncooked) nut is greyish brown in colour,
and when cut, shows reddish brown lines of albumen.
[ 187 ]
There are numerous varieties and many nuts of quality
are known by local names and fames all over India.
The betel palms are cultivated in pure-stand planta-
tions or in mixed gardens along with coconut palms,
bananas, oranges, limes, mangos, guavas, and numerous
other side plants of economic importance. It is a mari-
time palm, and requires a tropical climate for luxurious
growth. Its requirements are more or less the same
as in the case of the coconut, but it is very sensitive to
drought. In the tropics it grows well where there is a
heavy rainfall or under irrigation; it requires shade in its
early stages. It grows in a variety of soils from the lat-
erite soils of the west coast of India and Ceylon, through
Deccan along the east coast, to the alluvial soils of Ben-
gal and Assam, extending to Burma, Siam, Indo-China,
China and the islands of the North and South Pacific
Oceans.
Seedlings are raised in shady nurseries and transplanted
at an age of two or three years; seeds are also sown di-
rectly by dibbling the seeds in holes about 6-8 feet
apart. In Bengal, the first row of seeds is dibbled under
the shade of Hrythrina indica (the latter put in rows as
cuttings at 12-15 feet apart) and when the palms begin
to bear, the Erythrina plants are cut off and in their
place, and under the shade of the older palms, new rows
of seeds are dibbled. Bearing begins when the palms are
about 7-10 years old, but the plantation comes to full
bearing when it is about 15-20 years old and continues
to yield for from 80 to 60 years. The palms live over 100
years under good soil and cultivation.
b. Betel Vine: Piper Betle L.
The generic name Piper is probably a derivative from
the Sanskrit word Pippali for pepper. The name betel
(betle), meaning ‘‘simple leaf,’’ seems to come from betre
[ 188 ]
from the Malayalam language in South India. As in the
case of the betel palm, the betel vine has distinct names
in the various Indian languages. The betel vine, or pan
as it is sometimes called, is considered to be, probably,
anative of Java. Some even consider it to be indigenous
to South India, Ceylon and Malaya. Marco Polo, in his
travels, had seen betel leaves in Madras. Ibn Batula has
described both the betel nut and betel leaves; of the lat-
ter he writes: ‘‘a plant that grows like a grape vine. It
is trained over a trellis of canes like a vine or plants near
the coconut palm, where it clambers up like a vine or
pepper. It does not bear fruit; but it is the leaves which
are used and they are like those of the Bramble.’’ It is
extensively cultivated in India, Ceylon and Malaya.
Marco Polo mentions this leaf: ‘‘that people of India
have a habit of keeping in the mouth acertain leaf called
Tembul.’’ (The Sanskrit name is T'ambul; the Kannada
name J'ambula.) And he further comments: ‘‘that the
rich people and the King have these leaves prepared with
camphor and other aromatic spices, and quicklime.”’
Many other European travellers and others have noted
the use of betel leaf. Garcia da Orta has discussed the
use of pan in detail. ‘‘That Indians are in the habit of
keeping the nail of the right thumb pointed and sharp,
in order to remove the mid-rib of the leaf.’” The leaves
are used in the green and tender state.
The betel vine, a pepper, is an evergreen perennial,
twining on living supports—areca palms, species of /ry-
thrina, Sesbania, etc. It has a heart-shaped, simple, ex-
stipulate leaf, light green to deep green in colour, with
five to seven dorsally well marked veins.
Among the horticultural and agricultural plants which
need human care, the most cared for and the one which
is given the most delicate attention, is this vine. No
other plant seems to have so much tender and considerate
[ 189 ]
EXPLANATION OF THE ILLUSTRATION
Pirate XLVI. (Upper figure), Different types of
beeda, the commercial form of ready-made betel
chew. Left, northern India; center, Philippine Is-
lands; right, Siam.
From a painting by BLancue Ames.
(Lower figure). Map showing the area of cultiva-
tion of the betel palm, and the distribution of the
betel-chewing habit.
( 190 |
PiarE XLVI
Area where the Betel- nut palm is cultivated
Limit of use of Betel- nuts.
-
ee
attention by the cultivator. A worker in the vineyard
often reminds one of a servant in a palace, too careful to
spoil anything. For the successful growing of this vine,
one must need to have perfect knowledge of its require-
ments — adequate wind breaks, protective shade with
diffused light, perfect supports, rich and well-drained
humus soil, fairly constant temperature with a uniform
amount of moisture and regular manuring at intervals.
In some parts of India the vine is grown in specially
constructed houses of grass, reeds and mats, having flat
grass roofs so arranged as to admit diffused light.
As in the case of the betel nuts, there are numerous,
localized varieties of leaf; and in some details the culti-
vation varies from place to place. The vine is rarely
grown by the consumer, as it requires expert knowledge;
hence in India, the betel leaves travel long distances
from the cultivators to the consumers, i.e., to towns and
cities.
Harvest AND PREPARATIONS
a. Betel Nuts
The period from flowering to fruit ready to be har-
vested is nine to ten months. Flowering commonly oc-
curs throughout the year. The harvesting periods in
India begin in August and run up to the following Jan-
uary, the peak periods occurring mostly from October
to November.
For curing purposes, the fruits are collected just before
they are quite ripe. When one or two fruits in a bunch
begin to turn orange-yellow, the bunch is cut off and the
fruits are gathered. For raw nuts, all the fruits are al-
lowed to ripen to an orange-yellow colour.
The harvesting process is of interest. Special tree
climbers are employed for gathering the fruits. These
are the artisans of tree climbing. The climber uses a two
[ 191 |
to two and a half foot rope or banana fiber. This rope or
plantain fiber is tied together at the ends and twisted into
somewhat the shape of a figure eight in a horizontal po-
sition. The climber places a foot in each loop; the free
ends of his feet with his toes and the twisted middle por-
tion of the rope serve to clasp the cylindrical stem like
a clip. He first holds the stem with his hands, clasping
the stem at his chest level, and pulls his legs up with the
looped rope in his feet; at this level, he clasps the stem
with his feet and the rope, releases the hold of the hands
and pulls the whole length of the body up; then again
holds the stem with his hands, releases the hold of the
feet and pulls the legs up. These alternating holds and
pulls each time cover over two to three feet of the stem.
The climbers are so brisk and agile, that in seconds they
will be at the crown of the tree of sixty to eighty feet
in height. They go from tree to tree by swinging and
catching the next tree, as the areca palms are slender
and flexible, until the day’s harvest is completed and
without climbing down. It is easy to take a swing, if one
is bold enough, on an areca palm, but it is not possible
on coconut palms as they are not slender and flexible.
Once on the tree, the climber knows by experience which
branch to cut and which not to cut. He removes the
bunch and allows it to drop on the ground. The bunches
are then collected. Often if there is thick vegetation be-
low, the branches are caught in nets or lowered down by
means of a rope.
Semi-ripe fruits, that is, fruits to be used for curing,
are easy to husk. The pericarp is cut into halves without
bruising the seed and this is done by means of sickles or
special sharp blades fixed into a plank. One end of the
fruit is pushed against the blade, and when the blade
enters half way into the pericarp, and before it reaches
the seed, the fruit is pushed sideways which splits the
(192 J
pericarp into two parts and the seed drops out. This
shelling is done when the fruits are fresh and before they
dry up.
The curing process consists of boiling or cooking the
seeds. he shelled seeds are placed in copper or earthen
vessels with enough water to cover the seeds. For the first
charge, a small quantity of the previous year’s extrac-
tion, a decoction called chongaru, is added; or in the
absence of chongaru, a few pieces of the fresh bark of
Syzygium Cumini, Pterocarpus santalinus, Ficus religi-
osa, or other trees. Nobody seems to have considered
why these barks or chongaru are added to the first charge.
They may be likened to catalytic agents in chemical reac-
tions, and I suppose they are added to aid the tannin
reaction and the release of the same in the process of
cooking. In some cases a small quantity of sesame oil is
added. The charge is then heated to the boiling point.
The seeds are boiled until the tiny embryo drops out and
then they are removed by means of perforated ladles and
are dried for about a week. There are various modifica-
tions in this process of cooking. Unhusked fruits may be
boiled and then husked, sliced and dried; or the seeds
after husking may be cut into halves and cured, or sliced
into smaller pieces and then cured. Sometimes various
spices and jaggary are added during the boiling process
to increase the taste and aroma of the nuts. ‘‘Occasion-
ally the kernels are reboiled in milk to cater to fastidious
tastes. ”’
After three or four charges have been boiled in the
same decoction, the extract becomes semi-liquid; on
drying, this becomes a solid substance which is sold as
kath (cutch).
Seeds which are left whole during drying are called
unde-adike; the ones that are cut into cup-shaped halves
and dried are called battalu adike; those that are cut
[ 198 |
into smaller pieces and dried are called churu-adike, or
lavanga-churu. Thus the nuts are sold under different
names which vary according to whether they are uncut
whole nuts, or are cut into halves, or into smaller pieces.
This curing or cooking process is practiced throughout
India, particularly in Mysore and other parts of south
India.
The preparation of raw nuts in other nut-growing
countries is the same as in India.
Malaya produces four kinds of nuts: 1, whole nuts.
2, split nuts. 38, sliced nuts and 4, smoked nuts, and also
salted nuts in small quantities. Briefly some of the
methods followed in Malaya are as follows:
Pinang kossi. Ripe fruits are dried in heaps with oc-
casional turning for about three months before husking ;
after husking, the seeds are dried again for two or three
days before packing and marketing.
Pinang blah. Ripe truits are split into halves and dried
with the split sides uppermost for two or three days.
After the husk is removed, the seeds are dried again for
two or three days before marketing.
Pinang salai (smoked nuts). The smoking process is
done in a specially constructed mud oven with bamboo
gratings. The ripe fruits are placed on top of the bamboo
gratings and a fire is developed inside the oven with dry
betel-nut husks, the smoke of which escapes through the
mass of fruits; this smoke continues for about a week
before the fruits are taken out. They are then husked
and the seeds are dried for two to three days before
marketing.
Pinang asin. Semi-ripe green fruits are mixed with
salt and stored in sacks for two to three months before
husking. The fruits are then husked and the seeds dried
for two to three days before marketing.
[ 194 ]
b. Betel Leaf
Betel leaves are harvested throughout the year in ro-
tation, as the vine is evergreen and perennial. The leaves
are generally picked in the morning. They are picked
with at least one half inch of the petiole attached to the
leaf. Then these leaves are serially arranged one above
the other, the ventral surface of one lying on the dorsal
surface of the other and the petioles touching side by
side. Counting of the regularly arranged petioles is the
method adopted to count the number of leaves. The
leaves are counted in 20’s, 100’s and 500’s. In this ar-
rangement the leaves, alternating with banana or plan-
tain leaves, are packed into ball-shaped or barrel-shaped
large bundles, an operation which requires experienced
hands. Thus packed, they can withstand any amount of
handling and any distance of travelling. These artfully
packed leaves often travel long distances from the place
of cultivation to towns and cities, often on head loads,
on busses or in trains.
Besides supplying city and town markets, the betel
leaves are sold in the interior parts of India on a barter
system. Venders (usually women) carry leaves to each
village on alternate days or twice a week, and cater to
the village families and collect grains during harvesting
seasons.
OTHER INGREDIEN''S
a. Kath (cutch)
There are several types:
1. Kath from Acacia Catechu wood (pale catechu).
This is the crystalline substance prepared from a concen-
trated decoction of Areca Catechu wood. The chipped
wood of this plant is boiled in water to concentration
and the decoction is then cooled to get the crystalline
[ 195 ]
EXPLANATION OF THE ILLUSTRATION
Prare XLVI. Leaves of the betel pepper (Piper
Betle L.), the second of the principal ingredients
of betel chew, obtained in various Asiatic markets,
and showing variation in shape. A and D, Manila,
Philippine Islands; B and C, Bombay, India; E,
Bangkok, Siam.
PuateE XLVII
thick brown substance which is cut into blocks and sold
as kath by druggists. These cubes or blocks are redis-
solved in hot water and cooled into a semi-liquid for use
with pan; or the cubes are cut into small pieces and
used as such.
2. Kath, white kath, or gambier, from Uncaria Gam-
bir. The leaves and twigs are boiled in water and the
concentrated decoction is then cooled to get acrystalline
substance. This is cut into cubes or blocks and sold in
the market. The cubes are redissolved in water into a
semi-liquid for use with pan.
3. Kath from Areca Catechu seeds. As already de-
scribed under nut-curing, a semi-liquid is obtained by
boiling several charges of seeds. This chongaru, on dry-
ing, can be cut into blocks or cubes to be sold as kath.
b. Tobacco
There are various preparations of tobacco for use with
pan. ‘The tobacco used is a strong-smelling country-cured
tobacco with each leaf pressed into a sticklike form. It
comes to the market in small bundles.
1. Khaina. A thin layer of lime is applied to the left
palm and the chewing tobacco is rubbed and _ pressed
with the right-hand thumb which has also a thin layer
of lime. ‘This pressed lime-tobacco is then placed in the
mouth over the chewed pan. This limed tobacco is called
khaina.
2. Jharada. This is tobacco chewed with pan and other
ingredients and is used mostly in northern India.
3. Tombacu. In Hindustani, tombacu means tobacco.
The country-cured tobacco is powdered and moistened
with gur (jaggary) dissolved in water. It is variously
pressed and often is made into grain-like form for use in
[196 ]
the pan. It is frequently covered with gold or silver leaf,
and is the most expensive form of tobacco.
4. Kaddipudi. This is the cheapest form of tobacco.
It consists mainly of the stalks and the petioles of tobacco
plants. These sticky parts of the plants are crushed into
a coarse powder (/addipudi means powdered-sticks), pro-
cessed with jaggary and water, and pressed into bricks
or blocks and sold in the market.
5. Gundi. This is a mixture of country-cured tobacco,
coriander seeds and other spices. The tobacco is powdered
and fried; coriander seeds, and other spices, are each
fried until they have the correct smell and are then
coarsely powdered and mixed; finally the whole mixture
is scented with a resinous oil called chua; gundiis known
as kadapan in Orissa and Bengal.
c. Amber and lime
Amber. Amber is a fossil resin, but the term is also
loosely applied to many resins from living plants. It is
used to give an aromatic smell to pan.
Lime (quicklime). Originally lime for use in pan was
made from burned shells; but now it comes from burned
limestone; i.e., from calcium carbonate. Calcium ecar-
bonate is burned in special kilns, and is converted to cal-
cium oxide. On the addition of water, this calcium oxide
is changed to calcium hydroxide, which is known merely
as lime or quicklime, and is used in pan. Precaution is
always taken not to dry the lime completely in the open
atmosphere, as it will be partially reconverted to calcium
carbonate and become hard. Often the lime is variously
scented with camphor, or colored with saffron, and bot-
tled for sale. However, the plain lime is the form found
to be in most use. Each family buys its own stones and
prepares its own lime. The pan chewers carry their lime
[ 197 ]
in small metal containers or in small glass containers
called sunnadu-dabbhi (lime-container).
The purpose of using lime with the leaf is to neutra-
lize the burning and stinging acidity in the leaf. A leaf
taken without lime burns the mouth, and one gets a ter-
rible stinging sensation. That is why, knowingly or un-
knowingly, the pan chewers always use lime and smear
the leaf with it before placing it in the mouth. If an ex-
cess of lime is used, the effect would be the same, as the
lime will also burn the mouth. This is one precaution,
not to take a leaf without lime and not to use an excess
of lime, which the pan chewer knows by experience.
Generally the amount of lime applied to the leaf is very
small.
UTILIZATION OF Pan
a. Non-habitual chewing
In this type of chewing the pan is eaten entirely for
the carminative and salutary effect. Four different meth-
ods are used.
1. Pan trinity. Used either plain or with ingredients
(masala), this is the simplest form of pan. It varies de-
pending on a liking for astringency or sweetness. For as-
tringency, raw nuts are used; for sweetness, cured nuts.
The nuts are first placed in the mouth. While they are
softening, the leaves are cleaned, and the lime is applied
to them. They are then folded into small size and placed
in the mouth and chewed with the nuts which are already
in the mouth. The number of leaves used varies, but on
the average a person may use from three to five leaves
at a time. This form of chewing thus consists only of
betel nuts, betel leaves and lime. Some use various other
ingredients during the process of chewing; with cured
nuts, people who like the sweet taste may use pieces of
[ 198 ]
copra, cardamom seeds, cloves, nutmeg and mace; the
astringent loving ones use kath and cardamom seeds.
There is no hard and fast habit in these usages; one may
use only cardamom seeds with pan; or only cloves. They
often depend on the economic condition of the users, as
the ingredients, unlike pan, depend on the buying power
of the users.
Various kinds of pan boxes are used in northern India
to keep these pan trinities and their ingredients. These
are of various shapes, and often exquisitely ornated. They
are mostly made of brass and are divided into compart-
ments where the ingredients are placed. The boxes are
circular, with the compartments radiating, or rectangu-
lar; often they are two-storied, the upper story, which
has compartments, is fitted into the lower one; this up-
per one can be lifted out. The lower story, without com-
partments, contains the nuts and often a small nut slicer.
In southern India, the pan is offered on special plates;
in rich families on silver plates. The betel leaves are ar-
ranged on the plates, the nut mixture is placed ina silver
cup or in a glass bottle which is placed to one side, with
the lime in a small glass container or on a betel leaf; the
plate is then placed before the guests and relatives after
the meals.
2. Nut mixtures (pudi-adike). The nut mixtures also
differ in their ingredients depending on whether cured
nuts or raw nuts are used. Further, they differ in the
number of ingredients, aroma and freshness, depending
on whether they are prepared by the housewife for home
use or are commercial products, and also on the buying
capacity of the user.
Cured nut mutures. An average middle class well-to-
do family uses a nut mixture prepared in the following
way. Generally half nuts are broken into coarse pieces.
[ 199 ]
To these grated copra, grated nutmeg, crushed cardamom
seeds, broken pieces of cloves, broken mace, tiny pieces
of cardamom or cassia bark and saffron are added and
mixed well. The fresh grated copra gives a fatty coating
to the whole mixture and increases the aroma. Crystals
of sugar are then added. This mixture is always kept in
air-tight bottles to preserve its freshness. Occasionally
au very small quantity of camphor may be added to in-
crease the smell. ‘The same kind of mixture, sometimes
varying in ingredients, is prepared by small firms, or
restaurants, or pan shops for use in beedas; or for use
after a meal; often the mixture may further be scented
with amber or other resins. The mixture is placed in
small paper packages (each package contains a small
spoonful of mixture) and sold in the shops. Some restau-
rants keep these packages at the counters for the use of
the customers after meals.
Raw nut matures. The raw nuts are sliced, and fennel
seeds, caraway seeds, coriander seeds, cardamom seeds,
and broken cloves are added to them. The number and
kinds of seeds used vary very much in this mixture. In
northern India, hotels and restaurants very commonly
keep this mixture at the counter for the use of customers.
Many who may not like to chew the leaves use this
mixture.
3. Beeda. This is a commercial form of ready made
pan quid sold in the pan shops, in bazaars, restaurants
and hotels. There are two chief kinds of beedas, in many
ways different in their preparation and in their ingredi-
ents; one common to northern India and the other com-
mon to southern India.
Northern India. Each beeda contains at least two
leaves. The selected leaves are cleaned, the apex and
[ 200 ]
petioles are removed, and if the leaf is large, it is split
into two parts. The cleaned leaves are placed on the
palm of the left hand, one above the other with the dor-
sal surface upward; a dash of lime is applied to it by
means of a quill, and one or two dashes of kath, then
some pieces of sliced raw nuts are placed in the center;
a few seeds of cardamom are added, then the edges of
the leaves are folded over and a clove is stuck in to hold
the edges in place. It thus becomes a sort of inflated,
slightly irregular-looking square. The beedas are kept
arranged in trays with wet clothes to prevent drying off.
They are astringent and often bitter depending on the
amount of kath used. On account of the raw nuts and
the kath, if this pan is taken for the first time by one
unaccustomed to it, one may experience the choking
effect and all the other undesirable features attributed to
pan by writers. High grade beedas are often covered
with silver or gold leaf.
Southern India. In southern India, tender leaves are
selected for beeda, and each beeda contains at least two
leaves. The apex and petiole and to some extent the
mid-rib are removed, and a thin layer of one or two
spots of lime is applied to the dorsal surface. A spoonful
of nut mixture (consisting of cured half nuts broken into
small pieces and mixed with grated copra, grated nut-
meg, pieces of mace, crystals of sugar, cardamom seeds
and saffron) is then added to the leaves which are rolled
into a cone; the edges of the cone are pushed in and
stuck with a clove. These cone-like rolls are placed in
wire loops which are so arranged as to look like a green
flower bouquet; or are placed in plates arranged in a
pyramidal shape. These beedas are sweetish, aromatic
and tasty. Anyone can eat them whether or not for the
first time or by one unaccustomed to pan. There is no
[ 201 ]
astringency or bitterness in these quids. This is a very
common form of pan, chewed in southern India by non-
habituals, who eat it only for its taste and smell. These
are all commercial forms, but most families prepare their
own nut mixture and buy fresh leaves almost every day.
The home-made pan in southern India is something one
must taste to know what pan is really like.
4. Scented nuts. For this purpose only cured nuts are
used. Half nuts are scented with amber, camphor, resins
and other ground spices. These scented half nuts are each
packed in paper, like lemon drops or peppermints, to
keep the freshness of the nut. The half nuts are some-
times broken into two or three pieces and scented with a
ground mixture, as before, of amber, camphor, resins
and other spices, and to this, shelled melon seeds are
sometimes added.
b. Habitual chewing
In this form of chewing, indulged in for its own effect
or for the narcotic effect of the tobacco, mostly the lower
grades of nuts and leaves are used. In the majority of
cases, it is the poorer classes of people who have this habit.
For use with tobacco, the cured nut mixture, the southern
Indian beeda, and scented nuts are not utilized, not only
for economic reasons, but because they do not produce
any narcotic effects. The northern Indian beeda and raw
nuts and also their masala, however, are very well suited
for this kind of habitual chewing and for the use with to-
bacco. Ripe seeds and coarse leaves are better adapted,
since fibrous leaves and ripe seeds give a coarse base and
the bulk required to keep the tobacco in the mouth.
It is exactly the antithesis of the non-habitual chewing
where tender leaves and cured nuts are required, so that
they may be dissolved, so to say, in the mouth during
[ 202 ]
chewing. In this form of chewing, mostly pan trinity is
used with tobacco. Tobacco is also mixed with some
spices, as in gundi and jharada.
As in the plain pan chewing, the process is the same:
first the nuts are placed in the mouth, followed by leaves;
generally this class of chewers use more lime than the
non-habitual chewers. After the nuts and leaves are
chewed a little, they use the kind of tobacco to which
they are addicted ; some people prefer the rashed rhizome
of Smilax calophylla.
e. Ceremonial Utilization
To attempt to write about the ceremonial uses of
betel nut and betel leaves in India in this paper may be
likened to an attempt to write the cultural history of
the people of India in one sentence. There is no cere-
mony, there is no occasion, there is not a day in an In-
dian’s life where the pan does not play its part. The
happiest part of honeymoons, the happiest part of mar-
ried life is spent over pan. The young couples, on the
day’s work well done, retreating to their seclusion after
meals, talk out their hearts and pour out their affection
while the bashful young wife prepares pan quid with
selected leaves. The parents, in Indian families, plan out
and discuss the day’s outcome of work, the family affairs,
children, their education, over pan; while children clus-
ter about grandmother to hear fairy stories at leisure over
her pan.
Indian families, as a class, abhor anything narcotic,
anything alcoholic, and regard them as degrading to
the human soul. Even the pan, which has no narcotic
effect, is not allowed to be used by children or youths
at a tender age.
d. Medicinal Utilization
Betel nuts. Medically they are chiefly used in veteri-
[ 203 ]
nary practices as a vermifuge for tapeworms. Of all the
alkaloids in the nut, it is only arecoline which has toxic
properties. The juice of tender nuts is reported to act as
a laxative when given in small doses. This decoction, in
conjunction with other drugs, is used for costiveness and
dyspepsia. The burned nut is said to be used as a denti-
frice. A strong decoction of the ripe seeds is also used for
dyeing red and black shades.
Betel leaves. The essential oils from betel leaves are
used in treating catarrhal affections, inflammations of the
throat, larynx and bronchi; they exert an antiseptic ac-
tion, and also can be used with advantage in diphtheria.
The juice of fresh leaves is diluted and given as a dose
when the oils are not readily obtainable. The fresh juice,
mixed with lime, is applied to the throat in glandular in-
flammation. Many ayurvedic medicinal pills are admin-
istered with the fresh juice of betel leaves. The leaf. is
smeared with oil, warmed and applied as a poultice to
sores.
Economic IMporRTANCE OF THE PALM
The betel nut palm is one of the most important eco-
nomic plantation crops. The amount of nuts used in
India alone give some idea of its importance. It may be
safely said that about eight to ten million dollars worth
of nuts are used annually in India. She imports, besides
her own production, over 56,000 tons of betel nuts, and
absorbs almost all the surplus of the major betel-nut pro-
ducing countries like Ceylon, Straits Settlements, Indo-
nesia, ete.
As in the case of tobacco, coffee and tea, the betel nuts
are also taxed and the government gets considerable rev-
enue out of the nuts. There are areca nut merchant asso-
ciations in plantation areas to safeguard the plantation
interests and to regulate production and disposal of the
[ 204 ]
nuts. But there are a considerable number of small
patches of palms, grown near tanks and river sides for
family use or for catering to a group of villages, which
seldom come under statistics. Precise figures of acreage
under the palm are thus difficult to obtain. The statistics
regarding the actual acreage, total production, etc., are
incomplete; the figures often given in journals and books
must be taken with large margins. It is often noted that
the acreage for India and Burma is as follows:
Bombay State (Province) 22,700 acres
Madras - 108,600 ‘*
Mysore - 84,500 *‘
Ceylon 68,500 ‘
Burma 30,750 ‘‘
Bengal and Assam State (Province) greater acreage
than any other states (no definite figures).
The net profit per acre is not very high. It is stated
to be just over Rs. 100 per acre—an acre holds from
1,000 to 1,500 palms; each palm on the average yields
about 250-800 nuts; often the annual yield per acre is
up to 300 ewts. However, the yield of cured high-grade
nuts is much less; in Mysore, it is stated to be only 7
ewts. per acre. The prices vary very much according to
the grades from Rs. 50 to Rs. 8 per ewt.; the difference
of prices between first grade quality to inferior is often
as high as 5 to 1.
CONCLUSION
The use of pan in India has remained unchanged for
centuries (except for the impact of tobacco on pan as a
new form of chewing). Many of the conventional uses
of pan are remembered only through practice and are
handed down from parents to children. Pan is chiefly
used for chewing or for social purposes, rather than as a
[ 205 ]
narcotic or a drug. Of all the masticatories, fumatories
and liqueurs, the use of the pure form of pan is definitely
an innocent and harmless practice.
PLaNnts ASsocIATED WITH PAN
In each case the scientific name and family and the
part of the plant utilized is indicated, together with the
English and vernacular names.
i
a. Plants associated with chewing
Areca Catechu ZL. Palmaceae
Seeds (hemicellulose endosperm).
English: Betel nut, Areca nut; Sanskrit: Gur-
vaca; Hindustani and Persian: Swpari; Malayan
and Chinese: Pinang; Kannada, ete: Adike.
Piper Betle L. Piperaceae
Leaves.
English: Betel pepper, Betel leaves; Sanskrit:
Tambul; Hindustani: Pan.
Syzygium aromaticum (L.) Merr. & Perry
Myrtaceae
Dried unopened flower buds.
English: Clove; Sanskrit and Kannada: Lavang.
Elettaria Cardamomum (L.) Maton Zingi-
beraceae
Seeds.
English: Lesser Cardamom; Sanskrit: Upakun-
chica; Kannada: Hlakkt.
Amomum subulatum Povb. Zingiberaceae
Seeds.
English: Greater Cardamom.
Myristica fragrans Houtt. Myristicaceae
The seed and seed-cover (nutmeg and mace).
[206 ]
English: Nutmeg; Sanskrit: Jaiphal (Nutmeg);
Sanskrit: Jatri (Mace).
7. Cocos nucifera L. Palmaceae
Dried kernel (copra).
English: Coconut; Sanskrit: Nariyal; Kannada:
Tengu.
8. Cinnamomum zeylanicum Breyn. Lauraceae
Bark.
English: Cinnamon; Kannada: Darchini, Dal-
chini.
9. Cinnamomum Cassia (Nees) Nees ex Blume
Lauraceae
Bark.
10. Cinnamomum Camphora (L.) Nees & Eberm.
Lauraceae
Camphor obtained from the distillation of the
wood and leaves.
English: Camphor; Kannada: Karpura.
11. Dryobalanops aromatica Gaertn. f. Dip-
terocarpaceae
Camphor obtained from the distillation of the
wood.
English: Borneo Camphor.
12. Aquilaria Agallocha Rob. Thymelaeaceae
Resins.
13. Crocus sativus L. Iridaceae
Dried stigmas and styles of the pistil. Fairly com-
mon in use.
English: Saffron; Sanskrit: Kesar; Kannada:
Kesari, Kumkumkesari.
14. Pimpinella Anisum L. Umbelliferae
Dried fruits.
[ 207 ]
16.
17.
18.
19.
20.
21.
22.
23.
English: Anise; Sanskrit: Muhuri; Kannada:
Dodda-jirige.
. Foeniculum vulgare Miller Umbelliferae
Dried fruits.
English: Fennel; Hindustani: Pan-muhori;
Kannada: Saunf, Sonp.
Carum Carvi L. Umbelliferae
Dried fruits.
English: Caraway; Hindustani: Mita zerah.
Carum Bulbocastanum Koch Umbelliferae
Dried fruits.
English: Black caraway; Hindustani: Shah-zerah
Cuminum Cyminum L. Umbelliferae
Dried fruits.
English: Cumin; Sanskrit: Zira, jira; Kannada:
Jirige.
Nigella sativa L. Ranunculaceae
Dried fruits.
English: Black cumin; Hindustani: Kala Zerah;
Kannada: Kariirige.
Anethum graveolens L. Umbelliferae
Dried fruits.
English: Dill; Sanskrit: Misreys, Satapushpa.
Coriandrum sativum L. Umbelliferae
Dried fruits.
English: Coriander; Hindustani: Dhanya; Kan-
nada: Kothamburi.
Smilax calophylla Wail. Smilacaceae
Rhizome.
Tamarindus indica L. Leguminosae
Young leaves (rarely used).
[ 208 |
24.
26.
27.
28.
29,
30.
English: Vamarind; Hindustani: Ami; Kan-
nada: Hunise.
Uncaria Gambir (Hunt.) Roxb. Rubiaceae
W ood extract, Kath.
English: White cutch, Gambier.
. Acacia Catechu Willd. Leguminosae
Wood extract, Kath
English: Catechu; Hindustani: Kath; Kannada:
Kachu.
Saccharum officinarum L. Gramineae
Sugar from juice (stem).
English: Sugar cane; Hindustani: Ganna; Kan-
nada: Kabbu.
Beta vulgaris L. Chenopodiaceae
Sugar from juice.
English: Sugar beet.
Cucumis Melo L. Cucurbitaceae
Shelled seeds.
English: Melon.
Nicotiana Tabacum L. Solanaceae
Leaves and stalk.
English: Vobacco; Hindustani: Tombacu; Kan-
nada: Hogesoppu.
Nicotiana rustica L. Solanaceae
Leaves and stalk.
b. Plants associated with the preparation of pan
. Syzygium Cumini (L.) Skeels Myrtaceae
Fresh bark; used in the curing of betel nuts.
English: Black plum; Hindustani: Jambul;
Kannada: Nerale.
[ 209 ]
ho
Pterocarpus santalinus L. Leguminosae
Fresh bark; used in the curing of betel nuts.
English: Red sandalwood; Bengali: Rukto-
chandan; Hindustani: Chandan; Kannada: Gan-
dham, Kempugandha chekke.
Ficus religiosa L. Moraceae
Fresh bark; used in the curing of betel nuts.
English: Pipal; Sanskrit: Asvattha; Kannada:
Aswat, Aralt.
Sesamum indicum ZL. Pedaliaceae
Oil from seeds; used in the curing of betel nuts.
English: Sesame; Hindustani: 7%/; Kannada:
Allu.
Saccharum officinarum L. Gramineae
Gur (jaggary) made from juice; Gur is used in
the curing of betel nuts, and in several kinds of chew-
ing tobacco preparations.
c. Plants associated with the cultivation of pan
Sesbania aculeata Pers. Leguminosae
‘Tree serves as a supporter for the betel vine.
Bengali: Jayanti, Jiluga.
Sesbania grandiflora Pers. Leguminosae
Tree serves as a supporter for the betel vine and
as a shade tree.
Kannada: A gase.
Sesbania aegyptica Pers. Leguminosae
Serves for support and shade for the betel vine.
Hindustani: Saora.
Moringa oleifera Lam. Moringaceae
Serves for support and shade for the betel vine.
Kannada: Nugeze.
[ 210 |
Or
ep
Erythrina indica Lam. Leguminosae
Serves for support and shade for the betel vine.
Hindustani: Pangra.
Boswellia serrata Rob. Burseraceae
Stems used as poles to support the betel vine.
Hindustani: Sadlei.
Euphorbia Tirucalli Z. Euphorbiaceae
Used for windbreaks and as a fence for the vine-
yard.
Kannada: Kolu kalli.
Protium sp. Burseraceae
Used for windbreaks and as a fence for the vine-
yard.
Kannada: Konda mavu.
d. Plants grown with pan
All are grown for their economic importance either
intermingled or as a fringe plant in areca palm and betel
pepper gardens or plantations.
.
2.
Or
Cocos nucifera L. Palmaceae
Musa paradisiaca L. Musaceae
English: Plantain; Hindustani: Kela; Kannada:
Bale.
Musa paradisiaca L. subsp. sapientum (Z.)
O. Kize. Musaceae
English: Banana
. Mangifera indica L. Anacardiaceae
English: Mango; Kannada: Mavu.
Citrus aurantifolia (Chr.) Swg/. Rutaceae
English: Lime; Hindustani: Mimbu.
[ 211 ]
9.
10.
11.
13.
Citrus medica L. Rutaceae
English: Citron.
Citrus Limon (L.) Burm. Rutaceae
English: Lemon.
Citrus grandis (L.) Osbeck Rutaceae
English: Pomelo; Kannada: Chakkota.
Citrus sinensis (L.) Osbeck Rutaceae
English: Orange.
Syzygium Jambos (L.) Alston Myrtaceae
English: Rose apple; Kannada: Pannerale.
Psidium Guajava L. Myrtaceae
English: Guava; Kannada: Cheape.
Punica Granatum L. Punicaceae
English: Pomegranate; Kannada: Dalimbre.
Michelia Champaca L. Magnoliaceae
English and Hindustani: Champaca; Kannada:
Sampig.
14. Artocarpus heterophyllus Lam. Moraceae
English: Jackfruit; Kannada: Halasu.
15. Artocarpus altilis (Park.) Fosberg Moraceae
English: Breadfruitt.
16. Anacardium occidentale L. Anacardiaceae
English: Cashew; Kannada: Geru, Godumbi.
e. Plants used as catch-crops
17. Zingiber officinale Posc. Zingiberaceae
English: Ginger; Kannada: Shunt.
18. Curcuma longa L. Zingiberaceae
English: V'urmeric; Kannada: Harisina.
[ 212 ]
19. Manihot esculenta Crantz Euphorbiaceae
English: Cassava; Kannada: Maragenasu.
20. Ruta graveolens L. Rutaceae
English: Ruwe.
Many other economic plants of short duration are also
grown.
[ 218 ]
BIBLIOGRAPHY
Balfour, E. G.: Cyclopaedia of India and of eastern and southern
Asia, Madras, 1871.
Beccari, O.: The Palms of the Philippine Islands. Philipp. Journ.
Sci. 14 (1919) 295.
Blatter, E.S. J.: The Palms of British India and Ceylon. London,
1926,
Brown, W. H. and Merrill, E. D.: Philippine Palms and Palm Pro-
ducts. Philippine Bureau of Forestry Bull. no. 18. Manila, 1919.
Burkhill, I. H.: A Dictionary of the Economic Products of the Malay
Peninsula. London, 1935.
Candolle, A. de: Origin of Cultivated Plants. London, 1884,
Christy, T.: New Commercial Plants and Drugs. No. 5. London,
1882.
Elliot, H. M.: The History of India. Vol. 6. London, 1875.
Forster, E. M.: Abinger Harvest. London, 1922.
Garcia da Orta: Colloquies on the Simples and Drugs of India. Lon-
don, 1913.
Grist, D. H.: The Betel-nut Industry. Malayan Agric. Journ. 14
(1926) 219.
Hawks, E.: Pioneers of Plant Study. London, 1928.
Hill, A. F.: Economic Botany. New York, 1937.
Lewin, L.: Phantastica. London. 1931.
Lunt, W.: The Betel-nut Palm. Trinidad Roy. Bot. Gard. Bull.
Mise. Inf. 2 (1896) 204.
Martius, C. F. P. von: Historia Naturalis Palmarum. Leipzig, 1823-
1850.
Ridley, H. N.: Flora of the Malay Peninsula, London, 1925.
Sands: W. N.: Observations on the Betel-nut Palm (Areca Catechu
L.) and Betel Nuts. Malayan Agric. Journ. 14 (1882) 202.
Watt, G.: The Commercial Products of India. London, 1908.
The Wealth of India. A Dictionary of Indian Raw Materials and
Products. Vol. 1. New Delhi, 1948.
[ 214 ]
BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY
CamBripGr, MassacuuseTts, Marcu 18, 1951
VoL. 14, No. 9
NEW ORCHIDS FROM AFRICA
Be
V. S. SUMMERHAYES
(Royal Botanic Gardens, Kew)
THIS PAPER contains descriptions of two new genera
of Angraecoid orchids and new species belonging to vari-
ous other genera of African orchids. The type specimens
are in the Kew Herbarium unless stated otherwise.
Habenaria (§ Multipartitae) falciloba Swmmer-
hayes sp. nov. inter omnes species sectionis labelli lobis
lateralibus integris faleatim incurvatis, brachiis stigmati-
feris e basi liberis facile distinguitur.
Herba verisimiliter terrestris, circiter 1 m. alta, fere
omnino glabra; tubera non visa. Caulis erectus, robus-
tus, inferne circiter 1 em. diametro, fere teres, basi rad-
ices numerosas flexuosas 1-2 mm. diametro emittens,
inferne cataphyllis paucis vaginantibus apice obtusis su-
perioribus foliaceis instructus, superne circiter 9-foliatus,
Folia erecto-patentia vel suberecta, late lanceolata, apice
+ acuminata, usque ad 20 cm. longa et 6 cm. lata, sur-
sum decrescentia in bracteas abeuntia, tenuiter coriacea
vel subcarnosa. Racemus rectus, circiter 25 cm. longus
et 8 em. diametro, subdense circiter 35-florus; rhachis
teres, 3-4 mm. diametro; bracteae foliaceae, anguste
lanceolatae, acuminatae, subtus leviter carinatae, usque
ad 5 cm. longae et 1.5 mm. latae, inferiores flores ae-
quantes. Flores adscendentes; pedicellus cum ovario
[ 215 |
2-2.5 em. longus. Sepalum intermedium erectum, con-
vexum, oblongo-lanceolatum, apice acutum, circiter 2.75
em. longum et 8 mm. latum, dorso carinatum, 5-nervium;
sepala lateralia erecto-patentia, oblique lateque lanceo-
lata, leviter curvata, apice acuta, circiter 2.75 cm. longa
et 1 em. lata, 5—6-nervia, dorso carinata. Petala cum
sepalo intermedio agglutinata, falcata, e basi angusta
late lanceolato-cultrata, 2.5 cm. longa, supra medium 7
mm. lata, nervis superne+ divergentibus reticulatis. La-
bellum horizontaliter porrectum, ex ungue 6-7 mm. longo
dilatatum, tripartitum, in toto circiter 8 cm. longum et
1.5 cm. latum; partitio intermedia = inflexa vel incur-
vata, linearis, apice obtusa, 16 mm. longa, 2 mm. lata,
subcarnosa; partitiones laterales incurvatim falcatae, in-
tegrae, quam intermedia duplo latiores, apice acutae,
1.25-1.5 cm. longae, inferne 8.75-4 mm. latae, pluri-
nerviae; calcar infra labellum porrectum, leviter recur-
vatum, dimidio apicali clavato-inflatum, 2 em. longum,
superne 8.5 mm. diametro. Anthera 7 mm. alta, hippo-
crepimorpha; connectivus 5 mm. latus, medio 5.5 mm.
altus ; loculi fere horizontales vel leviter reclinati, 8.5 mm.
longi; canales + erecti, graciles, 4 mm. longi; stamino-
dia oblonga, parva, sessilia. Brachia stigmatifera basi
libera, crasse cylindrica, apice truncatim capitata, 9 mm.
longa, inferne pubescentia; rostelli lobus intermedius
triangulari-lanceolatus, acutus, 1.5 mm. longus, lobi
laterales cum anthera adnati, 1 cm. longi.
Bretaian Concao: Lualaba District, Kando, 60 miles W. of Jadot-
ville, March 28rd—April 4th 1931, De Witte 195 (Tyrer in Herb.
Brussels).
I have placed the species described above in sect. Mul-
tipartitae, since it agrees with this section in the general
habit, the large flowers, the veining of the sepals, the
character of the petals and the long slender anther canals,
as well as in the broad horseshoe-shaped anther connec-
[ 216 ]
tive. On the other hand, the side lobes of the lip are quite
entire and in this respect differ from all other species of
the section. Furthermore, the stigmas, instead of being
adnate to the lower side of the rostellar arms, are quite
free from the base and in general structure resemble these
organs in other sections of the genus. The anther con-
nective, though very broad, is comparatively tall, agree-
ing in this feature with Hi. egregia Summerh., but the
loculi are almost horizontal in position.
It is evident that in #7. falciloba we have a very aber-
rant species which may eventually have to be placed in
a distinct section of the genus.
Habenaria (§ Diphyllae) Keayi Summerhayes sp.
nov. affinis H. pilosae Schltr., a qua statura majore, folio
majore ovato vel late ovato apice acutiore, inflorescentia
13-30-flora, petali partitione postica prope basin dente
instructa, labelli caleari breviore = recurvato, anthera
distincte apiculata, staminodiis majoribus, brachiis stig-
matiferis lateraliter compressis satis distincta.
Herba terrestris, 40-50 cm. alta; tubera ellipsoidea
vel cylindrico-ellipsoidea, 1.5-4 em. longa, 5-15 mm.
diametro, breviter tomentosa; radices flexuosae, 1.5-2
mm. diametro, tomentosae. F’olium singulum vel rarius
unum multo minus additum, basale, humistratum, ova-
tum, vel late ovatum, leviter angulatum, apice acutum,
basi cordatum += amplexicaule, 8.5—-9.5 cm. longum, 6.5—
11 cm. latum, supra pilis longiusculis albis sparse indu-
tum, subtus glabrum, marginibus dense ciliatis, pallide
viride fusco-viridi-lineatum. Scapus erectus, teres, basi
usque ad 4 mm. diametro, cataphyllis numerosis lanceo-
latis vel lineari-lanceolatis acuminatis + adpressis infirma
rarius foliacea lanceolata usque ad 8 cm. longa instructus,
pilis debilibus albidis = dense indutus; racemus 12-19
cm. longus, 3-4 cm. diametro, laxiuscule 13—80-florus:
Co17
bracteae lanceolatae, acuminatae, pilosae, ovario pedicel-
lato breviores. lores erecto-patentes, pallide virides;
pedicellus cum ovario 10-12 mm. longus, dense pilosus.
Sepalum intermedium erectum, elliptico-ovatum, apice
obtusum vel fere rotundatum, circiter 7 mm. longum et
4 mm. latum, valde convexum; sepala lateralia deflexa,
oblique et leviter curvatim oblongo-lanceolata, apice sub-
rotundata, 7.5-8 mm. longa, circiter 38.25 mm. lata;
omnia sepala trinervia, dorso leviter carinata et pilis pa-
tentibus longiusculis instructa. Peta/a fere ad basin bi-
partita, glabra; partitio postica erecta, faleatim ligulata,
sepalo intermedio aequilonga, circiter 1.5 mm. lata, mar-
gine postico prope basin dente triangulari vel lineari in-
structa; partitio antica incurvatim vel falcatim adscen-
dens, circiter 1 cm. longa, basi circiter 0.75 mm. lata,
sursum sensim angustata, superne subcarnosa. Labellum
profunde tripartitum; pars basalis indivisa, 1 mm. longa
et 1.5 mm. lata; partitio intermedia fere recta, lineari-
ligulata, apice obtusa, marginibus valde recurvatis, car-
nosa, 9-10 mm. longa, vix 1 mm. lata; partitiones lat-
erales dimidio inferiore patentes, superne recurvatim
adscendentes vel circinatae, subcarnosae, circiter 138 mm.
longae, inferne 0.7 mm. latae, superne sensim angusta-
tae; calcar leviter recurvatim sigmoideum, dimidio api-
cali clavatim inflatum, 13-14 mm. longum, 1-1.5 mm.
diametro. Anthera erecta, 8—5.5 mm. alta, apice dis-
tincte apiculata; canales porrecti, circiter 1.5 mm. longi;
staminodia breviter stipitata, oblongo-elliptica, circiter
1mm. longa. Brachia stigmatifera antherae canalibus
breviora, carnosa, flabellatim capitata, = lateraliter com-
pressa; rostelli lobus intermedius erectus, magnus, an-
therae antipositus et eam aequans, elongato-triangularis,
acutus, antice excavatus.
Niaerta: Oyo Province, Ibadan Fuel Plantation, Sept. 1949, Keay
Forest Herb. 25395 (Type).
[ 218 |
This interesting species is clearly closely related to the
more southern A. pilosa Schltr. which occurs in North-
ern Rhodesia, the Katanga Province of the Belgian
Congo and the southernmost part of Tanganyika Terri-
tory. It is, however, a taller plant with larger differently
shaped leaves, while there are a number of less obvious
differences which are mentioned in the diagnosis. The
nearest relative in West Africa is H. Lecardi Kraenzl.,
but this has a very much longer spur. The present spe-
cies was described from dried and spirit specimens.
Vanilla polylepis Suwmmerhayes sp. nov. a V. im-
periah Kraenzl. foliis minoribus, bracteis vix 1 em. lon-
gis, floribus minoribus, labelli disco squamis + retrorsis
fimbriatis instructo satis distinguenda.
Herba alte scandens vel repens, succulenta, foliosa,
glaberrima; caules flexuosi, carnosi, usque ad 6 m. longi
vel ultra, usque ad 2 cm. diametro, ramosi, laxe foliati,
+ longitudinaliter plurisulcati, e nodis radicantes. Folia
oblanceolata, elliptico-lanceolata vel oblongo-lanceolata,
rarius elliptica vel oblongo-elliptica, saepius + curvata,
basi + rotundata, conduplicata et breviter petiolata vel
subsessilia, apice acutata vel leviter acuminata, 6-21 cm.
longa, 2.5-8 cm. lata; petiolus usque ad 5 mm. longus.
Inflorescentiae simpliciter racemosae, ramos laterales
terminantes, usque ad 7 cm. longae, dense usque ad 25-
30-florae ; rhachis carnosa, circiter 1 em. diametro; brac-
teae ovato-triangulares, acutae, subpatentes, 6-7 mm.
longae. Flores pallide virides vel viridi-albi; pedicelli
cum ovariis 4-6 cm. longi. Sepalwm intermedium oblan-
ceolatum, acutum, 8.5-5.25 cm. longum, 8.5-9.5 mm.
latum, 9-11l-nervium; sepala lateralia leviter oblique
oblanceolata, intermedio aequilonga, 9-12.5 mm. lata,
10-12-nervia; omnia sepala nervis Juxta apicem conver-
gentibus. Petala leviter oblique oblanceolata, acuta,
{ 219 ]
8.75-5.5 cm. longa, 10-14 mm. lata, 11—138-nervia, ner-
vis Juxta apicem vix convergentibus tribus centralibus
approximatis, dorso carina humili e basi per 2.5—4.5 cm.
longa in apiculo acuto terminante instructa. Labellum e
basi angustissima superne sensim sed valde dilatatum,
obovato-flabellatum, antice = obtuse incisum, 4.5—-5.5
em. longum, superne explanatum 8-8.5 em. latum, cum
columna per 2—2.75 cm. connatum, basi tubum angustum
superne dilatatum formans, antice marginibus valde un-
dulatis, multinervium; discus medio secus nervos tres
medianos squamis numerosis retrorsis instructus; squa-
mae anticae lineares, integrae, denticulatae vel dentatae,
intermediae transversae, ovatae vel oblongae, valde pec-
tinatae vel pectinato-incisae, posticae breviores, minus
pectinatae, postremae minutae papillis similes. Columna
fere recta, 2.75-4.25 cm. longa, semiteres, apice leviter
incurvata utrinque ala recurvatim oblongo-ovata obtusa
instructa; anthera quadrata antice angustata; medio
longitudinaliter sulecata; rostellum deflexum, ovato-
triangulare, circiter 2 mm. longum. Fructus immaturus
carnosus, cylindricus, sursum angustatus, bisuleatus, 11-
12 cm. longus, inferne 1.5 cm. diametro.
AnGoLa: Moxico District, island in river just above Lusavo Falls,
in evergreen vegetation, on ground and climbing up trees, Jan. 21st
1938, Milne-Redhead 4265.
Norruern Ruopesta: Mwinilunga District, River Matonchi, above
dam, in evergreen fringing forest, climbing up tree trunks and trail-
ing on the ground, Oct. 23rd—24th 19387, Milne-Redhead 2930 (Tyrr).
“*Stem and leaves deep green; one flower only out at time in any in-
florescence; sepals and petals greenish white; lip in front white, in
throat sulphur-yellow; column whitish; very faint mauve lines just
in front on channel inside lip; flowers when fading scented of vanilla. ’’
Mkusi District, Chiwefwe, 60 miles E.N.E. of Kapiri Mposhi, 1200
m. alt., probably from evergreen fringing bush, Nov. 1934, T'rapnell
1653, ““A very tall succulent liane; flowers greenish.’’ Abercorn Dis-
trict, Upper Lunzula River falls, 1650 m. alt., in evergreen riverain
forest, locally common, May 28th 1936, Burtt 6338 “‘A dark green
succulent climber ascending to 6 m.”’
[ 220 ]
Beiatan Conco: Katanga District, Elisabethville, 1440 m. alt.,
Nov. 22nd 1911, Rogers 10148. Same locality, in thick woods near
water, Sept. 1923, Von Hirschberg 28. ‘‘ Sepals and petals pale green;
lip snowy white shaded with orange to pale yellow, two mauve lines
from base to edge.’”’
This is apparently not an uncommon plant in the
fringing forests along the rivers and streams in a wide
area from eastern Angola to the Abercorn district of
Northern Rhodesia. As in its nearest ally, V. imperialis
Kraenzl., the lip and column, which are united nearly
to the top of the latter, form a trumpet-shaped tube
which is very narrow at the base and widens gradually
upwards. The sides of the lip are folded over the apex
of the column, but the lip is not distinctly lobed though
there may be a sinus of varying depth in the centre of
the apex. The scales forming the central crest are very
variable in character, the front ones being long and nar-
row, entire or more or less toothed, and resembling those
forming the crest of V’. imperialis. The central ones,
however, are broad and transversely placed, usually long-
pectinate or incised all around, but showing some ap-
proach to the simpler scales in V. africana Lindl., V.
ramosa Rolfe and similar species. Farther back the scales
are shorter but still pectinate or with long irregular
teeth, while in the lower part the outgrowths consist
entirely of short papillae-like hairs. The specific epithet
is given in allusion to this assemblage of scale-like out-
growths.
Malaxis (§ Katochilos) katangensis Swummer-
hayes sp. nov. affinis M. pygmaeae Summerh., a qua
statura majore, foliis plus duplo majoribus, sepalis uni-
nerviis, labello ambitu quadrato basi angustato distin-
guenda.
Herba terrestris, 9-12 cm. alta; tubera ellipsoidea,
circiter 1 em. longa et 5 mm. diametro. Caulis brevis,
[ 221 ]
erectus, 1-2 cm. altus, basi radices flexuosas graciles to-
mentosas emittens, superne cataphyllis paucis laxe vag-
inantibus apice+recurvatis acutis omnino vestitus, apice
2-3-foliatus. Folia patentia vel leviter recurvata, fere
humistrata, sessilia, late ovata vel elliptico-ovata, basi
caulem amplectentia, apice subacuta vel obtusa, 2.5—5
em. longa, 1.75-8 cm. lata, siccitate textura tenuiter
papyracea, plurinervia. Inflorescentia erecta, 8-10 cm.
longa, superne subdense multiflora; pedunculus 5-6 cm.
longus, gracilis, inferne fere teres, superne pluri-angula-
tus et angustissime alatus, cataphyllis 8-5 dissitis lanceo-
latis acuminatis 8-4 mm. longis instructus; rhachis pluri-
angulata; bracteae adpressae, lanceolato-ovatae, valde
acuminatae, 2.5-3.5 mm. longae, 1.5-2.25 mm. latae,
uninerviae. Fores pro sectione parvi, rhachidi+ adpressi,
virides, = hyalini; pedicellus cum ovario 2—2.5 mm.
longus. Sepa/um intermedium erectum, elliptico-lanceo-
latum, basi = rotundatum, apice subacutum vel obtu-
sum, 2—2.5 mm. longum, 1-1.25 mm. latum; sepala
lateralia libera, incurvatim deflexa, oblique ovato-lanceo-
lata, apice fere acuta, 1.75-2.25 mm. longa, 1.2-1.5
mm. lata. Peta/a patentia, leviter recurvata, lanceolato-
ligulata, apice obtusa, inferne leviter ampliata, 1.75—2.25
mm. longa, 0.8-0.6 mm. lata, marginibus integris vel
brevissime papilloso-ciliolatis; omnia tepala uninervia.
Labellum inferum, sessile, ambitu + quadratum, antice
rotundatum leviter emarginatum, basin versus sensim
angustatum, basi auriculis semiovatis 0.25-0.45 mm.
longis columnam amplectentibus instructum, in toto
(auriculis inclusis) 1.75-2.25 mm. longum, 1.6-1.75 mm.
latum, 5-nervium, basi tuberculis duobus humilibus,
medio pulvino dense puberulento quadrato instructum.
Columna incurvata, 0.7 mm. longa; anthera hemisphae-
rica; rostellum truncatum.
Betaran Conco: Katanga Province, Noots Farm, near Elisabeth-
[ 222 ]
ville, close to roots of large shady trees, very abundant, December
9th 1923, C. W. von Hirschberg 152 (Tyrer); Kisenga Valley, groups
of 20-100 plants growing at foot of large trees, Dec. 27th 1924, Von
Hirschberg 152 (Herb. Bruxell.); Kantu, by water, Jan. 26th 1908,
Kassner 2401.
This species is very similar to M. pygmaea Summeth.
in floral structure and general growth, but is a much
larger plant with larger leaves and minor differences in
the flower structure as indicated in the diagnosis. From
most other species of section Katochilos, it may be dis-
tinguished by the very short stem, the leaves thus appear-
ing almost on the ground. It appears to be decidedly
local in occurrence, though it may be very abundant
where it does occur.
Malaxis (§ Katochilos) pygmaea Summerhayes
sp. nov. inter species sectionis statura nana, cauli brevis-
simo, foliis et floribus parvis, labelli basi bituberculata
facile distinguenda.
Herba terrestris, pusilla, usque ad 5 cm. alta; tubera
1-3, ovoidea vel ellipsoidea, 6-10 mm. longa, 8-5 mm.
diametro, alba, cataphyllorum cicatricibus annulatis no-
tata; caulis erectus, 1-1.5 cm. longus, 1.5-—2 mm. dia-
metro, maxima pro parte subterraneus, cataphyllis pluri-
bus+omnino vestitus, apice 2—8-foliatus. Folia patentia,
fere humistrata, sessilia, late ovata vel elliptico-ovata,
basi caulem amplectentia, apice acuta apiculato-acumi-
nata, 7-20 mm. longa, 6-17 mm. lata, plicata, pluri-
nervia. Inflorescentia erecta, leviter flexuosa vel curvata,
1-4 cm. longa, superne dense multiflora; pedunculus
usque ad 2 cm. longus, teres vel obscure angulatus, gra-
cilis, cataphyllis paucis late lanceolatis caudato-acumina-
tis usque ad 3.5 mm. longis instructus; rhachis angulata;
bracteae + adpressae, lanceolato-ovatae, longe acumina-
tae, usque ad 2.5 mm. longae, uninerviae. Fores pro sec-
tione parvi, suberecti, rhachidi + adpressi, flavo-virides ;
[ 223 ]
pedicellus cum ovario circiter 2 mm. longus. Sepalwm
intermedium erectum, ovato-oblongum, basi rotunda-
tum, apice obtusum vel subrotundatum, 1.75-2 mm.
longum, circiter 1 mm. latum, 2-8-nervium; sepala
lateralia deflexa, incurvatim oblique ovata, apice subro-
tundata, 1.25-1.5 mm. longa, vix 1 mm. lata, 2- vel
subtrinervia. Petala patentia vel=recurvata, leviter ob-
lique anguste ligulata, apice obtusa, 1.25-1.5 mm. longa,
circiter 0.8 mm. lata, marginibus apice et basi exceptis
papillato-ciliolatis, uninervia. Label/um inferum (vel anti-
cum), sessile, ambitu ellipticum vel elliptico-quadratum,
basi in auriculas rotundatas columnam + amplectentes
decurrens, in toto (auriculis inclusis) circiter 1.75 mm.
longum, medio circiter 1.5 mm. latum, marginibus un-
dulatis, apice rotundatum leviter emarginatum, 5-ner-
vium, basi tuberculis vel callis humilibus orbicularibus
medio pulvino puberulento trapeziformi instructum.
Columna brevis, porrecta, subteres, apice truncata, cir-
citer 0.85 mm. alta; anthera hemisphaerica; fovea stig-
matica elliptica; rostellum truncatum.
NortHern Ruopesta: Mwinilunga District, just north of Mwini-
lunga on shallow soil overlying laterite in Brachystegia-Isoberlinia
woodland, Jan. 26th 1938, Milne-Redhead 4359 (Tyre) ‘“Tubers white,
ringed; leaves green, plicate; flowers yellow-green.”’
The species is described from over 25 specimens dried
or in liquid preservative. MM. pygmaea, as the name sug-
gests, is the smallest species of the genus found so far in
Africa, reaching at the maximum a height of 5 em. and
being usually much smaller. Another striking feature is
the very short stem, most of which is below the surface
of the ground, the leaves consequently being practically
at ground level.
The species in general floral structure belongs clearly
to the section Katochilos Schltr. in which the lip is an-
terior in position. This section includes six other species
in Tropical Africa and one in Madagascar.
[ 224 ]
Malaxis Hirschbergii Swmmerhayes sp. nov. affinis
M. Maclaudi (Finet) Summerh., a qua floribus majori-
bus, labelli marginibus basalibus incrassatis valde promi-
nentibus, labelli dimidio antico subito dilatato satis
distincta.
Herba terrestris, 10-15 cm. (fide collectoris 25 cm.)
alta, glaberrima; tubera non visa. Caulis erectus, circiter
5 cm. altus vel ultra, basi radices graciles flexuosas emit-
tens, superne cataphyllis 8-4 inferne vaginantibus apice
subfoliaceis liberis acutis vestitus, apice 2—3-foliatus.
Folia= patentia, 1-2 cm. petiolata, lanceolata vel ellip-
tico-lanceolata, apice acuta, in toto 8-7 cm. longa, 1.5—
3.3 cm. lata, siccitate molliter papyracea, 5-nervia. Jn-
Jlorescentia erecta, circiter 7-8 cm. longa, apice dense
multiflora; pedunculus 5-7 cm. longus, basi fere teres,
superne anguste bialatus, cataphyllis circiter 8 dissitis
lanceolatis valde vel caudato-acuminatis 6-8 mm. longis
instructus; rhachis brevis, fere 1 cm. longa; bracteae
lanceolatae, acuminatae, usque ad 6 mm. longae, ovario
pedicellato breviores. J/ores erecto-patentes, pseudo-
umbellati, pallide rubro-purpurei; pedicellus cum ovario
usque ad 13 mm. longus. Sepa/uwm intermedium oblongo-
lanceolatum, acutum, 5.5—5.75 mm. longum, 2—2.5 mm.
latum ; sepala lateralia libera, oblique lanceolato-oblonga,
subacuta, 5 mm. longa, 2—2.4.mm. lata. Peta/a elliptico-
lanceolata, apice acuta, 4.5-4.8 mm. longa, 1.75—-2.25
mm. lata, marginibus ciliolatis; omnia tepala trinervia.
Labellum inferum, sessile, ambitu late flabellatum, di-
midio basali marginibus incrassatis, in toto 4-4.5 mm.
longum, antice 4.7—4.9 mm. latum, dimidio apicali subito
dilatatum, transverse ellipticum, marginibus_ breviter
laceratis, basi callis duobus ligulatis humilibus parallelis
antice sensim dilatatis, callo dentiformi porrecto vel re-
curvato anteposito instructum, medio nervis tribus paral-
lelis utrinque nervo arcuato apice convergenti extus
[ 225 ]
ramoso instructum. Colwmna leviter incurvata, 1.5-1.75
mm. longa, semiteres; anthera semiellipsoidea, 0.5-0.6
mm. longa; rostellum ligulatum, anguste truncatum.
Betaian Coneo: Katanga Province, Kinsenga River, near Elisa-
bethville, in wet dark places at river’s edge under trees, only found
in this locality, Nov. 80th 1923, C. W. von Hirschberg 111 (Tyre;
Isorypr in Herb. Brussels).
This is clearly an ally of M. Maclaudu (Finet) Sum-
merh. and M. Chevalieri Summerh., both of which are
found in the general region stretching from French
Guinea to the Sudan along the north side of the Equa-
torial Forest belt. These are all characterised by the very
short apparently umbellate raceme, the relatively broad
fan-shaped lip and the tepals all more or less similar to
one another. The present species may be distinguished
most readily by the great development of the thickened
margin of the lower part of the lip which in this species
extends for half the total length; above this thickened
margin the lip suddenly widens to form a thin trans-
versely elliptical portion which is shortly lacerate all
round. In M. Maclaudi the basal thickened portion is
very poorly developed, while M. Chevalier lacks the
central projecting tooth-like callus.
Liparis odontochilos Swmmerhayes sp. nov. afh-
nis L. Deistelii Schltr., sed labello medio subito dilatato
marginibus lateralibus grosse dentatis differt.
Herba terrestris vel epiphytica, usque ad 13 cm. alta.
Caulis primarius repens, gracilis, teres, 1-2 mm. diame-
tro, cataphyllis arcte vaginantibus obtusis + obtectus,
radices flexuosas hie illic emittens; caulis secundarius
erectus, basi valde incrassatus, carnosus, pseudobulbum
anguste conicum formans, sursum sensim angustatus,
inferne cataphyllis subfoliaceis acutis vestitus, superne
3-4-foliatus, usque ad 5 cm. altus. Folia lanceolata vel
[ 226 |
elliptico-lanceolata, acuta, basi in vaginam angustata sed
vix petiolata, usque ad 8 cm. longa et 2.5 cm. lata, pli-
cata, textura tenuia, marginibus leviter undulatis. Jn-
Hlorescentia recta vel leviter curvata, usque ad 8 cm.
longa, superne sublaxe 2-10-flora; pedunculus teres,
angustissime alatus, 2-5 em. longus, cataphyllis paucis
lanceolatis acutis instructus vel nudus; rhachis angulata
vel anguste alata; bracteae erecto-patentes, lanceolatae,
acutae, usque ad 8 mm. longae. Fores erecto-patentes,
virides vel purpureo-striati vel purpureo-tincti; pedicel-
lus cum ovario circiter 8 mm. longus, gracilis. Sepalum
intermedium recurvatum, lanceolato-ligulatum, apice
acutum, basi auriculatum + amplexicaule, 4.5-8.75 mm.
longum, medio 1.25—2 mm. latum; sepala lateralia primo
fere ad apicem connata,synsepalum orbiculari-obovatum,
apice rotundatum, totum 3.5-6.25 mm. longum, 3.25-—
6.4mm. latum formantia, deémum=libera; omnia sepala
trinervia. Petala deflexa, linearia, apice obtusa, basi cum
sepalis adnata dilatata, 5-10 mm. longa, 0.35-1 mm.
lata, uni- vel subbinervia. Label/um e basi angusta brevi-
ter auriculata subito dilatatum, subflabellatim transverse
oblongum, antice truncatum, late emarginatum cum
apiculo interjecto, marginibus lateralibus grosse denta-
tis, 8-6.5 mm. longum, 4-7.5 mm. latum, basi callo
transverso obtuse bidentato instructum. Colwmna incur-
vata, semiteres, apice dilatata, late alata, 2.3-5 mm.
longa; anthera cucullata, antice emarginata.
Betc1an Conco: Ruwenzori, Lanuri, 1700-1800 m. alt., ona rotten
trunk in the shade, June Ist 1914, Bequaert 4629.
Uaanpa: Kigezi, impenetrable forest, creeping in moss on shady
bank in forest, Nov. 1948, Eggeling 5851.
Kenya Cotony: East Aberdare Mountains, Ocotea forest, in moss
on tree-fern stems, Dale 1860 (Typr); same locality, Thiririki River,
Nov. 1937, Cunningham-van Someren 21.
TANGANYIKA TERRITORY: Lossogonoi, S. by W. of Moshi, summit
of mountain, 2090 m. alt., in leaf mould on rocky ledge, coll. H.F.I.
Elliott, cult. and flowered Amani, May 20th 1944, Moreau 770.
| 227 |
This small creeping species is clearly related to L.
Deistel Schltr. which also occurs in Kast Africa. It may
be distinguished by the lip being suddenly broadened in
the front part with the outer margins very coarsely
toothed. The basal callus is more clearly transverse, with
two lateral rounded knob-like teeth.
Bulbophyllum encephalodes Swummerhayes sp.
nov. affine B. oreonasti Rehb.f. et B. unifoliato De
Wildem., ab illo pseudobulbis unifoliatis, pedunculo
quam rhachide duplo longiore, ab hoc pseudobulbis quad-
rangularibus, foliis latioribus, floribus majoribus + atro-
rubris nec flavis, ab utroque labelli dimidio apicali rimis
complicatis sinuosis verrucisque ita instructo ut cerebrum
simulet satis differt.
Herba epiphytica, glaberrima. Rhizoma repens, usque
ad 35 cm. longum, 2-3.5 mm. diametro, primo vaginis
scariosis 5-7 mm. longis, 5-12 mm. distantibus instruc-
tum, sub pseudobulbis radices caespitosas flexuosas ra-
mosas tenues griseas emittens. Pseudobulbi 1. 5-8 (saepius
4-6) cm. distantes, elliptico-oblongi vel conico-elliptici,
+ compressi, acute tetragoni, apice + truncati, mono-
phylli. Folia oblonga, elliptico-oblonga vel elliptica, basi
conduplicata, in petiolum brevissimum angustata, apice
rotundata, 3-12 cm. longa, 1.2-8.2 cm. lata, subcarnosa,
coriacea, supra leviter canaliculata. Scapus erectus, racemo
+ recurvato vel dependente, 13-26 cm. altus; peduncu-
lus rectus vel + leviter curvatus, teres, 11-24 cm. longus,
1-2 mm. diametro, vaginis 4-7 arctissimis, obtusis, dis-
tantibus, 5-13 mm. longis instructus; rhachis valde com-
pressa, 2-7 cm. longa, 3 mm. lata, marginibus leviter
sinuatis, dense multiflora. Fores distichi, e rhachidis
marginibus alternatim exorientes, sessiles, rubro-virides
vel + atrorubri; bracteae patentes vel leviter recurvatae,
rigidae, ovatae vel lanceolato-ovatae, acuminatae, 2.5—5
[ 228 ]
mm. longae, scariosae. Sepalum intermedium lanceolato-
ovatum, acuminatum, 4-5 mm. longum, 2.5—-8.25 mm.
latum, valde convexum, + incurvatum; sepala lateralia
oblique lanceolato-ovata, longiuscule subulato-acumi-
nata, 4.5-5.25 mm. longa, basi 2.25-3 mm. lata, mento
brevi 0.75—-1 mm. longo; omnia sepala trinervia. Petala
lanceolato- vel elliptico-oblonga, acuta, 2.5—3 mm. longa,
0.6-1 mm. lata, uninervia. Labellum valde carnosum,
ambitu ovoideo-ellipsoideum, leviter curvatum, totum
2.5-8 mm. longum, 1.5-1.75 mm. latum, inferne carinis
humilibus + parallelis, medio praesertim subtus rimis
complicatis sinuosis reticulatis + transversis, apice verru-
cis numerosis irregularibus instructum. Columna brevis,
semi-teres, 2 mm. alta, stelidiis brevibus acutis, andro-
clinio leviter excavato; anthera + hemisphaerica, leviter
retusa; ovarium crassum, 2-8 mm. longum, teres, levi-
ter plurisuleatum. Capsu/ae ellipsoideae, 7 mm. longae,
4—5 mm. diametro.
Ueanpva: Ankole, Kalinzu Forest, Aug. 1936, Eggeling 3196;
“*Epiphyte, flowers greenish with purple spots and claret centre.’’
Kenya Cotony: Kericho District, Dec. 1933, Gray 3; Kakamega
Forest, 1500 m. alt., bank of river in bush country, Oct. 1939,
Tweedie 492; ‘“Epiphyte, flowers plum-coloured with darker streaks. ”’
TANGANYIKA Territory: East Usambara Mts., Bomole, near Amani,
1050 m. alt., growing on quinine and other trees in comparatively
dry situations, Nov. 29th 1940, Moreau 6 (Tyre); ““Tepals mainly
green, edged with maroon, upper side of labellum dark maroon.’’
Between Amaniand Monga, 900 m. alt., epiphytic in topmost branches
of evergreen rain-forest of Cephalosphaera, Tylostemon kuao and Pari-
nari holstii, Nov. 18th 1936, Greenway 4742; ‘‘ Flowers greenish
crimson,”
Betatan Conao: Katanga, near Elisabethville, on trees near stream,
July 31st 1930, Hutchinson 4122; ‘‘Flowers deep crimson.’’ Lubun-
bashi, epiphytic in gallery forest, Nov. 1927, Quarré 908; Katanga,
no locality, epiphyte in forest plantation, Feb. 1935, Quarré 4454.
NortTuern Ruopesia: Mwinilunga District, River Mwanamitawa,
about 12 miles west of River Lunga, epiphytic on tree trunk in Cryp-
tosepalum woodland, Aug. 22nd 1930, Milne-Redhead 945; “‘Flowers
reddish-purple, lip dark purplish-black,column pinkish,anther white. ’’
Same district, slope east of River Matonchi and by river, on trees in
[ 229 ]
Brachystegia woodland and also in evergreen vegetation, flowered May
16th 1938, Milne-Redhead 4375; ‘‘Sepals green, mottled maroon,
petals greenish-yellow, edged maroon, lip white, side lobes maroon,
anther lemon-yellow. ’’
This is clearly a widely spread species, mainly in sa-
vanna vegetation but also in rain-forest. Its affinities and
characteristic features are set out in the diagnosis. The
frequently more or less reflexed or almost pendulous ra-
ceme is an interesting character. The lip is a truly re-
markable structure which is very difficult to describe. It
is very fleshy, more or less ellipsoidal, and somewhat
curved especially in the lower part. Here there are two
low parallel keels which run forward for about two-thirds
of the lip becoming lower and more undulate or sinuous
towards their apices. The body of the lip is smooth at
the base, but in the centre the surface becomes increas-
ingly broken up by very curved sinuous fissures forming
a coarse and irregular reticulation. In the apical third
this fissuring becomes so pronounced as to produce a
series of large irregular warts covering the whole surface.
The general effect is reminiscent of a brain, hence the
specific epithet. These irregularities are well shown in
several gatherings in liquid preservative and are not due
to drying.
Bulbophyllum porphyrostachys Swmmerhayes
sp. nov. affine B. lupulino Lindl. a quo rhachide medio
anguste nuda, bracteis purpureis apice breviter incurva-
tis, labello magis carnoso alis angustissimis apice verru-
coso facile distinguendum.
Herba epiphytica, glabra; rhizoma repens, teres, 2-8
mm. diametro, basi pseudobulborum radices numerosas
flexuosas ramosas emittens. Pseudobulbi 2.5—4 em. dis-
tantes, erecti, saepius 4-angulati, ellipsoidei vel oblongo-
ellipsoidei, 2-4 cm. longi, 1.5—2 cm. diametro, basi cata-
phyllis paucis, scariosis apiculatis + vestiti, apice diphylli.
[ 280 ]
Folia erecto-patentia, e petiolo brevi ligulata vel oblan-
ceolato-ligulata, apice rotundata vel apiculata, 8-18 em.
longa, supra medium 1.8—-2.3 cm. lata, saturate viridia,
supra nitentia, secus medium leviter canaliculata, subtus
leviter carinata. Scapus e basi pseudobulbi ortus, rectus
vel leviter curvatus, folia aequans vel brevior, usque ad
15 cm. altus; pedunculus 4-6 cm. longus, teres, viridis,
cataphyllis 5-8, laxe ochreatis, acutis, 7-10 mm. longis
fere omnino vestitus; rhachis anceps, leviter fractiflexa,
5-10 cm. longa, 2.5-8 mm. lata, viridis; bracteae disti-
chae, patentes, imbricatae, ovato-cymbiformes sed valde
lateraliter compressae, apice leviter incurvatae, 9-18 mm.
longae, 5—6 mm. latae, fusco-purpureae, glabrae. Flores
sessiles, quam bracteae multo breviores, purpurei vel
virides purpureo-maculati, semiclausi. Sepaluwm interme-
dium incurvatum, valde concavum, triangulari-ovatum,
apice acutum, apiculatum leviter cucullatum, circiter 4
mm. longum, 1.8—-2 mm. latum; sepala lateralia oblique
ovata, apice faleatim decurvata, acuminata, fere 4 mm.
longa, circiter 2.5 mm. lata; omnia sepala subcarnosa,
viridia, intus purpureo-notata. Petala linearia, leviter
faleatim curvata, apice obtusa vel subacuta, 2.8 mm.
longa, 0.8 mm. lata, viridia vel purpurea. Labellum valde
carnosum, sectione semiteres, ambitu ellipticum, supra
leviter excavatum, medio canaliculatum, subtus valde
convexum, utrinque ala angustissima denticulata instruc-
tum, apice verrucosum, circiter 2 mm. longum, 1.6—1.7
mm. latum, saturate purpureum. Columna semiteres,
2.3 mm. alta, superne anguste alata, stelidiis ovatis acutis
antheram aequantibus; anthera cucullata, ambitu ellip-
tica, antice breviter producta; fovea stigmatica quadrata,
antice labio prominente instructa; Ovarium crassum, V1Xx
angulatum, deorsum angustatum, 2 mm. longum.
Nigeria: Benin Prov., Okomu Forest Reserve, near Nikrowa, in
high forest on crown branches of fallen Corynanthe, Dec. 1947, cult.
[ 231 ]
and flowered Ibadan, Oct. 1948, Keay in Forest Herbarium 22711
(Tyre); Benin Prov., Sapoba Forest Reserve, in broken high forest,
on crown branches of large Jrvingia gabunensis, Oct. 28th 1946, Keay
& Onochie in Forest Herbarium 21590.
This very striking species is clearly closely related to
B. lupulinum Lindl., possessing similar large spreading
imbricating bracts and a very similar flower structure.
It may, however, be easily distinguished by the different
shape of the quite glabrous purple or crimson bracts,
which here have a slightly incurved apex and conse-
quently a slight dip in the upper margin. In B. lupuli-
num the upper margin is either almost straight or evenly
convex all along, while the bracts are fawn coloured and
dotted all over with small furfuraceous hairs. In our
species the bases of insertion of the two rows of bracts
are separated so that there is a narrow median strip of
the rhachis visible nearly to the apex, whereas in B.
lupulinum the bases of the bracts often overlap in a ver-
tical direction so that the rhachis is only visible on elon-
gation. The lip in B. porphyrostachys is extremely fleshy
with a rounded convex lower surface and a flat or slightly
convex upper surface. At the edge of this are the very
narrow toothed wings which are a little wider in front.
The apex of the lip is furnished with a number of small
warty excrescences.
Triceratorhynchus Swmmerhayes gen. nov.
Epiphytica. Caulis brevissimus, plurifoliatus. Molia
linearia, apice vix bilobulata. IJnflorescentiae patentes,
elongatae, pluriflorae. Flores parvi, pedicellati. Sepala
et petala libera, inter se similia, patentia, anguste lan-
ceolata. Labellum calearatum, simplex, concavum, lan-
ceolatum, ecallosum, tepalis aequilongum; calear gracile,
ex ore angusto cylindricum, labelli laminam superans vel
aequans. Columna brevis, crassa, apice truncata; andro-
clinium leviter excavatum, dorsum versus reclinatum;
[ 282 ]
anthera postice hemisphaerica, antice in appendicem tri-
angularem obtusam eroso-ciliatam producta; pollinia 2,
ellipsoidea, stipitibus duobus brevibus paulo supra me-
dium viscidiorum exorientibus, viscidiis duobus distinctis
elliptico-ligulatis, stipitibus multo longioribus ; rostellum
utroque productum, viscidiis amotis utroque trifurcatum,
sursum lobis lateralibus quam intermedio satis longiori-
bus, deorsum lobis lateralibus triangulari-dentatis lobo
intermedio fere subulato longiore ; fovea stigmatica quad-
rata vel fere orbicularis.
Species unica: 7". viridifiorus Summerh.
This plant, which looks rather like an Angraecopsis or
small Angraecum, is chiefly characterised by the remark-
able structure of the rostellum. This is attached to the
apex of the column and is produced both upwards and
downwards from the point of attachment to form a
roughly ovate shield-like plate. On removal of the two
elongated viscidia the rostellum is left trifureate in both
directions, the upward prolongations being longer than
those downwards. The general effect is to produce a
tridentlike or 3-horned structure in both directions. The
two vertical erect lateral lobes are somewhat incurved
and nearly twice as long as the narrower very acute cen-
tral lobe; on the other hand, the downwardly produced
middle lobe, which is very similar to the upward one, is
considerably longer than the quite short lateral lobes.
Another interesting feature is in the anther which is
produced in front into a broadly triangular beak which
is erose-ciliate all around the front margins.
The genus is perhaps most closely allied to Sphyra-
rhynchus Mansf. which has a short column with the broad
rostellum produced both upwards and downwards to
form a hammer-head like structure. In the latter genus,
however, there is only a single viscidium and on removal
of this the rostellum is obtusely bilobed or almost entire.
[ 233 ]
Both genera are small plants with few-flowered racemes
and relatively small leaves, though in Sphyrarhynchus
these are much broader than in the present genus.
Triceratorhynchus viridiflorus Swmmerhayes sp.
nov.
Planta pusilla, epiphytica; caulis brevissimus, circiter
5 mm. longus, 38—5-foliatus, radices numerosas flexuosas
emittens. /olia suberecta vel + reflexa, linearia, saepius
+ curvata, apice obscure inaequaliter bilobulata vel in-
tegra, acuta, lobulo breviore dentiformi, lobulo longiore
usque ad 1.5 mm. longo, tota 1-5 cm. longa, 1-3.5 mm.
lata, supra = concava, carnosula. Inflorescentiae ex ax-
illis foliorum exortae, patentes vel horizontales, pauci-
vel pluriflorae, usque ad 6 em. longae; pedunculus usque
ad 2 cm. longus, gracilis, teres, vaginis paucis acutis
brevibus instructus; rhachis pedunculo similis; bracteae
breviter ochreatae, late ovatae, acutae, 1—1.5 mm. longae.
Flores usque ad 9, 4-6 mm. distantes, patentes, virides
ad flavi; pedicellus cum ovario 4-5 mm. longus, saepius
medio + geniculatus. Sepalum intermedium lineari-
lanceolatum, acutum, 4.25—-5.5 mm. longum, 0.75-1.25
mm. latum, trinervium; sepala lateralia oblique lanceo-
lato-ligulata, leviter curvata, subacuta, 5.25—6.25 mm.
longa, 0.9-1.2 mm. lata, tri- vel subquadrinervia. Petala
curvatim lineari-lanceolata, acuta, 4.25—5.25 mm. longa,
vix 1 mm. lata, trinervia; omnia tepala patentia vel +
recurvata. Labellum valde concavum, ambitu late lan-
ceolatum, acutum, medio obscure trilobatum, lobis later-
alibus erectis brevissimis rotundatis, lobo intermedio
superne incrassato recurvato, totum 4.75—-5.5 mm. lon-
gum, explanatum circiter 8.5 mm. latum, plurinervium ;
calear ex ore angusto cylindricum, leviter incurvatum,
apice obtusum, 5.5-8 mm. longum. Columna generis,
circiter 0.75 mm. longa; anthera circiter 1 mm. longa;
[ 234 ]
pollinia 0.25—-0.5 mm. longa, stipitibus + geniculatim
decurvatis apice subspathulato-dilatatis 0.2 mm. longis,
viscidiis ligulatis utroque rotundatis marginibus hyalinis
0.7 mm. longis; rostellum viscidils amotis sursum trifur-
catum, lobis lateralibus leviter convergentibus cornibus
similibus circiter 0.5 mm. longis, lobo intermedio tri-
angulari-subulato quam lateralibus satis breviore, deor-
sum breviter trifurcatum, lobis lateralibus brevibus tri-
angularibus acutis, lobo intermedio fere duplo longiore
subulato-lineari, circiter 1.5 mm. longo.
Ueanpa: Bunyoro, Budongo Forest, June 1935, Eggeling 2058;
same locality, epiphytic on Olea Welwitschii, cult. and flowered Busin-
giro, June-July 1944, Eggeling 5492 (Tyrer).
Kenya Cotony: Kakemega Forest, 1500 m. alt., June 1944, E.W.
Carroll; comm. F.M. Tweedie 648.
This plant is probably generally distributed in the for-
ests of this region, but it is easily overlooked on account
of its small size and inconspicuous green flowers.
Eggelingia Swmmerhayes gen. nov.
Epiphytica. Caulis monopodialis, elongatus, basi radi-
cans, superne laxe foliatus. Folia ligulata, apice bilobu-
lata. Inflorescentiae ex axillis foliorum ortae, brevissimae,
pauciflorae. Flores parvi, pedicellati. Sepala libera, +
patentia, lanceolata vel oblonga, aequalia. Petala ob-
longa, quam sepala paulo minora. Labellum calearatum,
simplex, ovatum, acutum, ecallosum, sepalis aequilon-
gum; calcar fere rectum, ore pro rata angustum, cylin-
dricum, prope apicem paulo dilatatum, labelli laminae
aequilongum. Columna brevis, crassa, apice truncata,
semiteres; androclinium leviter excavatum; anthera
hemisphaerica, antice in appendicem acutam producta;
pollinia 2, ovoidea vel subpyriformi-ovoidea, stipite lig-
ulato apice truncato dilatato = incurvato, viscidio rec-
tangulari-elliptico stipite paulo breviore, rostellum de-
[ 285 |
flexim porrectum, triangulare, viscidio amoto profunde
bifidum; fovea stigmatica rectangularis.
Species duae, Africae tropicae incolae.
Species typica: 7. igulfolia Summerh.
This small genus resembles very closely some species
of Tridactyle and, in vegetative characters, certain species
of Angraecum, all having rather long straggling stems
with spreading ligulate leaves. The very short inflores-
cences are also very reminiscent of 7V'ridactyle such as
T. anthomaniaca (Rehb.f.) Summerh. and 7\. Scottellit
(Rendle) Schltr., especially those species in which the lip
is almost entire. Mggelingia differs, however, in the quite
entire lip, which lacks the thickened basal auricles so char-
acteristic of T7idactyle, and in the very large rectangular
viscidium, which is nearly as long as the stipes and longer
than the pollinia. The genus is best placed next to 7'r-
dactyle.
The genus is distributed from the Gold Coast in the
west to the forests of western Uganda in East Africa,
being widely spread in the forests of the eastern Congo
Basin. It is named in honour of Dr. W. J. Eggeling
who has done so much towards increasing our knowledge
of the epiphytic orchids of Uganda.
Eggelingia ligulifolia Swmmerhayes sp. nov.
Planta epiphytica vel rarius saxicola. Caules saepius
singuli, pauciramosi, saepius = penduli vel e basi des-
cendente sursum arcuati, graciles, usque ad 45 cm. longi,
2-3 mm. diametro, basi vel inferne radices griseas flex-
uosas emittentes, foliorum delapsorum vaginis omnino
vestiti, superne laxiuscule foliati. Molia circiter 1 cm.
distantia; vagina arcta, leviter transverse rugulosa, sic-
citate longitudinaliter costulata, internodium aequans
vel paulo longior, apice laminae opposita ligula subulata
usque ad 7 mm. longa instructa; lamina sessilis, = patens
236 |
vel rarius recurvata, ligulata vel lineari-ligulata, apice in-
aequaliter bilobulata vel subacuta, lobulo longiore usque
ad 6 mm. longo, supra secus medium canaliculata, sub-
carnosa, 2-7 cm. longa, 5-8 mm. lata. Inflorescentiae ex
axillis foliorum ortae, brevissimae, 2—8-florae ; pedunculus
fere nullus; rhachis fractiflexa, usque ad 8 mm. longa;
bracteae triangulares, acutae, pedicello cum ovario duplo
breviores. Flores albi; pedicellus cum ovario 4.75—-6.75
mm. longus. Sepalum intermedium late lanceolatum,
acutum, 5.25-6.75 mm. longum, 2.2—2.6 mm. latum,
trinervium; sepala lateralia oblique lanceolata, leviter
curvata, acuta, 4.5-7.75 mm. longa, 2—2.3 mm. lata, 2-
nervia. Petala oblonga vel oblongo-lanceolata, subacuta
vel obtusa, leviter obliqua, 4.5—6 mm. longa, 1.75-2.25
mm. lata, subtrinervia. Labellum valde concavum, late
lanceolatum vel lanceolato-ovatum, apice leviter acumi-
natum, 4.5-5.75 mm. longum, explanatum 2.75-3.5
mm. latum, plurinervium; calcar fere rectum, cylindri-
cum, vix inflatum, 4-5 mm. longum. Columna crassa,
1.25-1.75 mm. longa, structura generis; pollinia 0.6—0.8
mm. longa; stipes 1-1.7 mm. longus; viscidium 1.3—1.4
mm. longum; ovarium leviter 6-sulcatum, pilis brevibus
furfuraceis instructum.
Beteran Coneo: Angi, lava plains, on lava, Jan. 21st 1914, Bequaert
5819.
Ueanna: Kigezi, Kayonza Forest, Ishasha Gorge, cult. & flowered
Busingiro, September 1944, Eggeling 5379; Kigezi, impenetrable
forest, Nov. 1948, Eggeling 5845; Toro, 1500 m. alt., on trees in
forest, coll. C. H. Lankester, cult. & flowered Kampala, Nov. 26th
1921, Snowden 736; Toro, Itwara Forest, Aug. 1943, Eggeling 5372
(Type).
This species occurs in the region around the main rift
valley in East Africa, between Ruwenzori and the Vir-
unga Mountains further south. Although usually a forest
plant, it also occurs on lava in the voleanic areas. This is
true of a number of other orchids which are normally
epiphytic.
[ 237 ]
Eggelingia clavata Swmmerhayes sp. nov. ab E.
hgulifola Summerh. foliis pro rata paulo latioribus, in-
florescentiis usque ad 6-floris, floribus minoribus, labelli
dimidio apicali clavatim inflato satis differt.
Planta epiphytica. Cau/es singuli vel caespitosi, sim-
plices, saepius + penduli vel e basi descendente sursum
arcuati, graciles, usque ad 35 cm. longi, 2-8 mm. dia-
metro, basi radices flexuosas ramosas emittentes, inferne
foliorum delapsorum vaginis fere omnino vestiti, superne
laxiuscule foliati. Jota circiter 1 cm. distantia; vagina
arcta, paulo anceps, leviter transverse rugulosa, interno-
dium aequans vel paulo brevior, apice laminae opposita
ligula subulata usque ad 7 mm. longa instructa; lamina
sessilis, patens vel + recurvata, ligulata vel oblongo-
ligulata, apice inaequaliter obtuseque bilobulata, lobulo
longiore usque ad 6 mm. longo, supra secus medium
canaliculata, subcarnosa, 2—5 cm. longa, 5-9 mm. lata.
Inflorescentiae ex axillis foliorum ortae, brevissimae, 2—
6-florae; pedunculus usque ad 38 mm. longus, cataphyllis
ochreatis 1-2 ovatis obtusis instructus; rhachis fracti-
flexa, usque ad 8 mm. longa; bracteae e basi breviter
vaginante triangulari-ovatae, subacutae, pedicello cum
ovario plus duplo breviores. Flores albi vel flavido-albi;
pedicellus cum ovario 8-4 mm. longus. Sepalum inter-
medium late lanceolatum, late oblongo- vel elliptico-
lanceolatum, acutum vel obtusum, 3-8.75 mm. longum,
1.6-1.9 mm. latum, trinervium; sepala lateralia oblique
lanceolata vel late lanceolata, apice acuta, 8.25—4.75 mm.
longa, 1.25-1.75 mm. lata, 2-3-nervia. Petala oblonga
vel oblongo-lanceolata, obtusa, leviter obliqua, 2.25-8.75
mm. longa, 1-1.75 mm. lata, 1—8-nervia. Labellum valde
concavum, late lanceolatum vel ovato-lanceolatum, levi-
ter acuminatum, 8-3.5 mm. longum, explanatum 2.25—
3.25 mm. latum, plurinervium; calcar fere rectum, ore
angustum, dimidio apicali clavato-inflatum, apice rotun-
[ 238 ]
datum, 2.5-4 mm. longum, 1—1.25 mm. diametro. Col-
umna crassa, 0.75—-1 mm. longa, structura generis; pol-
linia ovoidea, 0.4-0.5 mm. longa; stipes 1—-1.15 mm.
longus; viscidium circiter 1 mm. longum; ovarium levi-
ter 6-sulcatum, pilis brevibus furfuraceis instructum.
Gotp Coast: Ashanti, Ofinso, Aug. 1938, Cor 99.
Gazson: Upper Ngoumyé River, Lake Bandoungou, in forest, Dec.
30th 1924, Le Testu 5160 (Type).
Bete1an Conco: Between Weko and Bengamisa, N. of Stanleyville,
around sources of River Ngula, July 16th 1939, Louis 15484; Penghe,
banks of River Itur, in forest, Feb. 6th 1914, Bequaert 2334; Kibale
District between Irumu and Mambasa, in forest, Oct. 1931, Lebrun
4173; no locality, 1920, Lacomblez 565.
This species resembles very closely KH. ligulifolia
Summerh., from which it may be distinguished by the
characters given in the diagnosis, particularly the smaller
flowers and much swollen spur. It also differs in a num-
ber of other less obvious features which can be ascer-
tained by careful comparison of the two descriptions. It
is clearly a forest plant, so it will be interesting to see if
it turns up in the Cameroons and in the Southern Ni-
gerian forests.
[ 239 ]
A REVISION
OF THE GENUS ANGRAECOPSIS
BY
V. S. SUMMERHAYES
(Royal Botanic Gardens, Kew)
THE GENUS Angraecopsis was first proposed by Kraenz-
lin* in 1900 for a plant from Usambara in Tanganyika
Territory, namely A. tenerrima Kraenzl. The author
pointed out most of the distinctive characters of the
genus, particularly the unequal sepals and the adnation
of the laterals to the anticous part of the petals, though
he seemed a little confused over the morphological status
of the parts concerned. He also stated that there is a
single viscidium, but this is clearly the result of faulty
observation. Afterwards he appears to have rejected his
own genus since he redescribed A. tenerrima in 1914 as
a species of Angraecum.
In his comprehensive account of the Orchidaceae in
**Die Orchideen’’ published in 1914, Schlechter’ recog-
nises the genus and adds three species to the original
A. tenerrima. These were A. parviflora (Thou. ) Schltr.
from the Mascarene Islands, A. ischnopus (Schltr. )
Schltr. from West Africa and A. falcata (Thunb. ) Schltr.
from Japan. He says little about the characters of the
genus except to point out that the three-lobed lip sepa-
rates it easily from Aérangis Rehb. f.
A few years later he gives a more complete treatment
of the genus in his general revision of the Angraecoid
Orchids*. In this the number of species has risen to seven,
‘Engl. Bot. Jahrb. 28 (1900) 171.
? Die Orchideen (1914) 600-601.
® Beih. Bot. Centralbl. 36, Abt. 2 (1918) 139-141.
[ 240 ]
in spite of the fact that 4. falcata has, quite properly,
been removed to a separate genus, now known as Neo-
jinetia Hu. Three of the added species are from West
Africa, the fourth is from the Comoro Islands; all had
been described some time previously, but had not been
recognised as belonging to Angraecopsis.
The latest formal account of the genus is that of the
Madagascar species by Perrier de la Bathie*, in which
five species are enumerated of which three are additions.
All these species have distinct viscidia to the pollinia
in spite of any statements to the contrary, though in
dried specimens the two viscidia, which often lie side by
side, adhere to one another during drying and are difh-
cult to separate on later examination.
Recently several orchids, which seem referable to 4n-
graecopsis on general grounds, have been collected in
different parts of East Africa. These species, however,
all have a common viscidium to the two pollinia. It
seems most suitable to include them in a distinct section
of the genus.
Examination of all the above forms shows that, al-
though they exhibit considerable diversity in many re-
spects, there is a recognisable facies common to the whole
genus, not only as regards the vegetative parts, but also
in the floral structure. Characteristic external features
are the short stem with a few somewhat oblique curved
leaves, the relatively long slender inflorescences and the
rather small or very small somewhat spidery flowers. In
the latter the lateral sepals are characteristically longer
than the intermediate, often somewhat spathulate and
projecting together forwards in a parallel position. The
petals are triangular, usually with the anticous margin
dilated into a forward pointing lobe, the lower part of
this being adnate to the base of the lateral sepal; often
* Humbert. Fl. Madag. Orchid. 2 (1941) 80-86.
[ 241 ]
the very acute apex is sharply reflexed so that it is not
at first apparent. The lip is commonly trilobed but is
almost entire in a few species; the spur has a narrow
mouth, but is rather variable otherwise. The rather short
acute rostellum projects forward or slightly downward ;
it is apparently variously cleft after the removal of the
viscidia. It should be pointed out, however, that the
flowers are rather small and the state of preservation of
some of the species is not good enough to be certain of
the exact details of the rostellar structure.
As aresult of my investigations I now place the total
number of recognised species at 14, of which three are
newly described later in this paper. Since several of the
species are very small plants it seems probable that there
may yet be further similar species awaiting discovery ;
some of the mainland species have been gathered only
in the past 20 years or so.
The affinity of Angraecopsis appears to be with Mys-
tacidiwm, from which it differs in the unequal sepals, the
characteristic petals, the normally 8-lobed lip, the nar-
row mouth to the spur and the side lobes of the rostel-
lum not being papillose or pubescent.
Angraecopsis Kraenzlin in Engl. Bot. Jahrb. 28
(1900) 171—Schlechter, Die Orchideen (1914) 600-601,
et in Beih. Bot. Centralbl. 86, Abt. 2 (1918) 1839-141—
H. Perrier de la Bathie in Humbert, Fl. Madag. Orchid.
2 (1941) 80-86.
Herbae epiphyticae. Caules breves, usque ad 5 cm.
longi, radices flexuosas simplices vel ramosas emittentes,
superne paucifoliati. Folia disticha, saepius ligulata, sed
interdum oblonga vel linearia, saepe = curvata. Inflores-
centiae saepissime ex axillis foliorum delapsorum exortae,
singulae vel plures, racemosae, pauci- vel pluri- (rarius
multi-) florae, pedunculo saepius rhachide longiore.
[ 242 ]
Flores alternati, parvi vel rarius mediocres, albi virides
vel flavo-virides, longiuscule pedicellati; bracteae parvae,
basi vaginantes. Sepa/a inter se libera, lateralia quam
intermedium saepius longiora + spathulata et parallele
porrecta, interdum omnia sepala = patentia. Peta/a an-
tice basi cum sepalis lateralibus adnata, saepe oblique
triangularia apice reflexa. Labellum saepius medio vel
infra medium trilobatum vel tripartitum, rarissime integ-
rum, lobis lateralibus brevibus usque interdum interme-
dium superantibus; calear ex ore angusto cylindricum
vel apice + inflatum. Co/wmna brevis, crassa, apice trun-
cata; androclinium leviter excavatum; anthera + hem-
isphaerica, antice breviter producta; pollinia duo, sphae-
roidea vel ovoidea, stipitibus duobus linearibus vel rarius
superne dilatatis, viscidiis distinctis vel rarius viscidio
communi; stigma excavatum; rostellum breviter vel
rarius longiuscule productum, saepius porrectum, Vis-
cidiis amotis = 2—8- lobatum vel partitum.
Species typica: 4. tenerrima Kraenzlin.
Sectio 1. Hu-angraecopsis Summerhayes sect. nov.
Plantae parvae vel mediocres: pedunculus rhachidem
aequans vel saepius longior; sepala valde inaequalia; pe-
tala triangularia vel fere dolabriformia margine antico
valde dilatata, acuta; labellum trilobatum vel triparti-
tum, lobis lateralibus saepius linearibus vel ligulatis; pol-
liniorum stipites pro rata longi, viscidiis distinctis parvis.
Species 10, Africae tropicae et Insulae Mascarenarum
indigenae.
Species typica sectionis: A. tenerrima Kraenzlin.
Sectio 2. Cardiochilus Summerhayes sect. nov.
Planta pusilla; pedunculus rhachide saepius brevior;
rhachis fractiflexa; sepala inaequalia; petala elliptico-
ovata, rotundata; labellum integrum, basi leviter corda-
tum; polliniorum viscidia distincta.
[ 243 ]
Species unica sectionis, Africae tropicae indigena: A.
pusilla Summerhayes.
Sectio 8. Coenadenium Summerhayes sect. nov.
Plantae parvae; pedunculus rhachidem aequans vel
brevior; sepala + inaequalia; petala ovata vel lanceolata,
margine antico vix dilatata, subacuta; Jabellum triloba-
tum vel integrum, lobis lateralibus brevibus dentiformi-
bus; polliniorum viscidium commune, pro rata magnum,
stipites saepius geniculatim conniventes.
Species 8, Africae tropicae orientali indigenae.
Species typica sectionis: A. breviloba Summerhayes.
KEY TO SPECIES
Pollinia with 2 separate stipites and viscidia
Petals triangular or broadly triangular with the anticous margin
much dilated, adnate to the lateral sepals along the lower anti-
cous margin, apex acute usually sharply reflexed
Spur 2.5 cm. or more long; relatively large plants with ligulate
leaves 7-20 cm. in length; lip distinctly trilobed; petals
markedly triangular; lateral sepals + spathulate, much longer
than the dorsal sepal; peduncle long, flowers about 6-12,
spaced out rather laxly along the rhachis
Lateral lobes of lip from slightly shorter to slightly longer
than the middle lobe, broad and obscurely lobulate at the
apex; petals much broader than long; spur 5-6.5 cm. long;
leaves usually from 1.5-2.5 em. broad, narrower in small
individuals . . . . . . . . . . . . . .) tenerrima
Lateral lobes of lip much shorter and narrower than the mid-
dle lobe, quite entire; petals about as long as broad; spur
2.5-4.5 em. long; leaves usually 7-15 mm. broad, rarely
broader .........2...+... 8. gracillima
Spur less than 2 cm. long, usually much shorter
Lateral lobes of lip well developed, never less than half the
length of the middle lobe
Spur shorter than the lip, much swollen at the apex; lateral
lobes of lip about as long as or longer than the middle
lobe; petals slightly longer than broad, 1-nerved; lateral
[ 244 ]
sepals not markedly spathulate; small plants with leaves
up to 8 cm. long
Spur nearly 4 mm. long; rhachis about equalling the pe-
duncle, 6-17-flowered ; lateral sepals 4.5-5.5 mm. long;
leaves broadly ligulate, 6-10 mm. broad
7. dolabriformis
Spur about 2 mm. long; rhachis shorter than the pedun-
cle, 6-7-flowered; lateral sepals 3.5-4.5 mm. long;
leaves linear, 2-6 mm. broad . ... . . 8. tridens
Spur longer, often much longer, than the lip, usually not
much swollen at the apex
Petals broader than long (length measured from apex to
base of central vein)
Leaves small, elliptical-lanceolate or oblong, up to 5
em. long and 12 mm. broad; inflorescences 4—6 cm.
long; rhachis about as long as peduncle, about 5-7-
flowered ; lateral lobes of lip slightly longer than the
middle lobe; spur about twice as long as the lip, not
atallswollen . ....... =... 41. ischnopus
Leaves large, strap-shaped, 20-35 em. long, 2.5-3.25
em. broad; inflorescences at least 10 cm. long, rha-
chis much shorter than peduncle, 10—20-flowered ;
lateral lobes of lip distinctly shorter than middle
lobe; spur longer than the lip, swollen at the apex
4. macrophylla
Petals longer than broad
Leaves broadly strap-shaped or elliptical-oblong, scarce-
ly curved, 10-24 mm. broad; petals 3-nerved
Leaves elliptical-oblong, 4-10 em. long, 10-24 mm,
broad, with two slightly unequal rounded lobes at
the apex; inflorescences up to 15 em. long, rhachis
2-4.5 em. long, 7-13-flowered ; lip about 4.5 mm.
long, lateral lobes much shorter and narrower than
the middle lobe, spur 5-8 mm. long _ 5. elliptica
Leaves broadly strap-shaped, 9-15 cm. long, 11-22
mm. broad, unequally lobed at the apex; inflores-
cences 12-22 cm. long, rhachis 4—9 cm. long, 10-
25-flowered ; lip about 6.5 mm. long, lateral lobes
nearly as broad and as long as the middle lobe,
spur about 16 mm. long . . . . . . 6. trifurca
Leaves narrowly strap-shaped, usually somewhat
[ 245 ]
curved, up to 18 cm. long but only 4-12 mm. broad,
acute or very unequally lobed at the apex with the
shorter lobe usually tooth-like; petals 1-nerved ; in-
florescences up to 17 cm. long, rhachis short, 1-3.5
em. long, up to 12-flowered; lip less than 4 mm.
long, lateral lobes distinctly shorter than the middle
lobe, spur 6-9 mm.long . . . . . . 9. parviflora
Lateral lobes of lip very small, spreading ; flowers very small
(dorsal sepal 2 mm. long); lip more or less cruciform with
very short side lobes and thickened apex; spur longer
than the lip, much swollen at the apex; leaves ligulate,
2cm. long,4-7 mm. broad . . . . . . 10. Pobeguinii
Petals rounded ovate, anticous margin scarcely dilated, adnate to
lateral sepals for a short distance at the base; lip entire, broadly
ovate, truncate or slightly cordate at the base; spur slightly
longer than the lip; leaves linear, 1.5-5 em. long, 1.5-5 mm.
broad; rhachis + zigzag, flowers 5-10 mm. apart 11. pusilla
Pollinia with 2 separate stipites but a common viscidium; petals not
much differentiated from the dorsal sepal, only shortly adnate to
the lateral sepals
Lip quite entire or with very obscure rounded side lobes, broadly
ovate, spur about 2 cm. long; stipites of pollinia long and slen-
der, not geniculate, viscidium convex, ovate; leaves ligulate, up
to 9 em. long and 1 em. broad . . . . . . . . 12. holochila
Lip with distinct acute tooth-like side lobes about or below the mid-
dle, linear and very fleshy, spur 12 mm. long or less; stipites of
pollinia short and thick, geniculately connivent, viscidium ob-
long; dorsal sepal 3-4 mm. long; leaves very small, usually less
than 2.5 cm. long
Spur scarcely inflated in the apical part, 10-12 mm. long; side
lobes at or just below the middle of the lip 13. amaniensis
Spur markedly inflated in the apical part, about 4.5 mm. long;
side lobes well below the middle of the lip 14. breviloba
ENUMERATION OF SPECIES
Section 1. Mu-angraecopsis Summerhayes
This section contains all the species in which the char-
acteristic features of the genus are best developed. In
particular the long more or less spathulate forward-
[ 246 ]
projecting lateral sepals give the flower a distinct and
readily recognised facies. ‘The species vary from small
plants with a few leaves about 3-4 cm. long up to the
imposing 4A. macrophylla in which the strap-shaped leaves
reach a total length of 85 cm. These are very reminis-
cent of the leaves in certain species of Chamaeangis and
Diaphananthe.
1. Angraecopsis ischnopus (Sc//tr.) Schlechter,
Die Orchideen (1914) 601, et in Beih. Bot. Centralbl.
36, Abt. 2 (1918) 140— Summerhayes in Hutchinson
& Dalziel, Fl. West. Trop. Afr. 2 (1986) 462.
Angraecum ischnopus Schlechter in Notizbl. Bot.
Gart. Berlin 4 (1905) 170.
Sierra Leone: Loma Mountains, E. slope of Mt. Bintumane, Oct.
1944, Jaeger 289.
Frencuw Guinea: Nimba Mountains, in mountain forest, Sept. 1947,
Schnell 3729; same locality, Oct. 1947, Schnell 3851.
British Cameroons: Buea, Deistel (Tyrr).
This species is characterised by the broad petals, these
being considerably broader than long, and the rather long
slender spur. ‘The rhachis is about the same length as
the peduncle, the few flowers being spaced out at rela-
tively long intervals. The Sierra Leone specimen is in
fruit only, but appears to be correctly placed in this
species.
2. Angraecopsis tenerrima Kraenzlin in Engl.
Bot. Jahrb. 28 (1900) 171 — Schlechter in Beih. Bot.
Centralbl. 36, Abt. 2 (1918) 141.
Angraecum tenerrimum Schlechter in Engl. Bot.
Jahrb. 88 (1906) 164.
Angraecum amaniense Kraenzlin in Engl. Bot. Jahrb.
51 (1914) 396.
TANGANYIKA Territory; East Usambara Mountains, Ngwelo (Nque-
lo) Forest, June 1899, Scheffler; Amani, Braun 3211; Ndola, 960 m.
[ 247 ]
alt., evergreen forest, May 1950, Verdcourt § Greenway 216; Kwam-
kuyu Falls, 840 m. alt., on rock scarp by waterfall, Jan. 1931, Green-
way 2845; Mahenge Distr., Muhulu Mountains, 1200 m, alt., in mist
forest, Feb. 1932, Schlieben 1844.
This and the next species, 4. gracillima (Rolfe) Sum-
merh., may be distinguished from all other species by
the long spur, which reaches a length of 2.5—-6.5 em.
A. tenerrima has the longest spur in the genus, from 5—
6.5 cm. in length, and may also be distinguished from
A. gracillima by the nature of the lateral lobes of the
lip. In the present species these are as long as the middle
lobe and almost parallel-sided to the apex which is often
obscurely lobed or bluntly toothed, whereas in 4. gra-
cillima the lateral lobes are shorter, often much shorter,
than the middle lobe and quite entire at the apex.
3. Angraecopsis gracillima (Rolfe) Summerhayes
in Kew Bull. (1987) 465.
Mystacidium gracillimum Rolfe in Kew Bull. (1913)
144 — Schlechter in Beih. Bot. Centralbl. 36, Abt. 2
(1918) 126.
Betet1an Conoco: Between Masisi and Walikale, 1300 m. alt., in
transition forest, March 1932, Lebrun 5156; Kamatembe, Bishakishaki
River, N.W. of Mt. Nyamlagira, April 1934, De Witte 1516; Nyam-
lagira, Kanamaharagi Lava, 1600 m. alt., in sclerophyllous vegeta-
tion, Feb. 1945, Germain 3599; Goma, by Lake Kivu, in lava plains,
1650 m. alt., in forest, March 1948, Purseglove 2616.
Ucanpva: Toro District, 1800 m. alt., in forest, flowered Botanic
Gardens, Entebbe, May 1939, Chandler 2792; Ankole District, Lake
Lutoto, flowered Busingiro, Sept.—Oct. 1944, Eggeling 5470; Mengo
District, Mabisa Forest, July 1908, Brown 444 (? Iso-Tyern); Masaka
District, South Buddu, Fyffe 184; Mbale District, Mt. Elgon, Tracey
Falls, 1650 m., alt., July 1917, Snowden 505; Mt. Elgon, 1800 m. alt.,
in forest, June 1924, Snowden 902.
Kenya Cotony: North Kavirondo District, Kakamega-Kaimosi
Forests, 1500 m. alt., flowered Nairobi Sept. 1938, Cunningham—Van
Someran 47; Kakamega Forest, 1200 m. alt., April 1938, Van Someran
(Coryndon Museum no. 8411); same locality, 1650 m. alt., in forest,
Aug. 1942, Tweedie 590; South Kavirando District, Kisii, 1500 m.
[ 248 ]
alt., Napier (Coryndon Museum no. 6355); Kericho District, Sotik,
Kipsonoi River, 1500 m. alt., Gray; Chepalungu Forest, 1800 m.
alt., on dead Acacia in grassland, Honoré 2544.
Evidently widely spread in the East African forest
regions. The differences between this and ZL. tenerrima
Kraenzl. are given under that species and in the key.
The leaves are normally narrower in this species, but one
gathering from the Belgian Congo (Lebrun 6156) has
leaves as broad as those of /. tenerrima.
4. Angraecopsis macrophylla Summerhayes sp.
nov. affinis A. gracillimae (Rolfe) Summerh., a qua foliis
multo majoribus, inflorescentiis densifloris, calcari plus
duplo breviore apice inflato satis differt.
Herba epiphytica; caulis brevis, usque ad 5 cm. lon-
gus, decurvatus, inferne radices flexuosus ramosas verru-
cosas 1-2 mm. diametro emittens, superne paucifoliatus.
Folia omnia pendentia, ligulata, + curvata, basi leviter
angustata, apice valde inaequaliter bilobulata, lobulo
breviore brevissimo vel fere nullo, lobulo longiore usque
ad 2 cm. longo, obtuso vel subacuto, in toto 15-35 ecm.
longa, 1.5-8.5 ecm. lata, subearnosa, dorso leviter cari-
nata, pallide viridia; vagina 2-3 cm. longa, compressa,
siccitate striata. Inflorescentiae ex axillis foliorum delap-
sorum exortae, patentes vel = dependentes, circiter 15
cm. longae, simpliciter racemosae; pedunculus teres,
11-12 cm. longus, 1—-1.5 mm. diametro, vaginis circiter
5 dissitis arctis acutis dorso carinatis usque ad 5 mm.
longis instructus; rhachis quam pedunculus paulo cras-
sior, leviter fractiflexa, 2.5-3.5 cm. longa, dense usque
ad 20-flora; bracteae breviter ochreatae, triangulares,
acutae, 1-2 mm. longae. Flores patentes, de colore nil
constat; pedicellus cum ovario fere 1 cm. longus. Se-
palum intermedium erectum, late oblongo-ovatum, apice
rotundatum, convexum, circiter 2.5 mm. longum et 2
[ 249 ]
mm. latum; sepala lateralia subspathulato-ligulata, in-
curvata, parallele porrecta, circiter 5.5 mm. longa et
superne 1.75 mm. lata; omnia sepala trinervia. Peta/a
oblique triangularia, margine antico valde dilatata sepalo
laterali adnata, apice angustata, acuta, dimidio superiore
reflexa, 2.5 mm. longa, basi 2.8 mm. lata, subtrinervia.
Labellum basi utrinque angulatum, superne trilobatum,
5.5 mm. longum; lobus intermedius oblongus, sursum
sensim angustatus, obtusus, 2.5 mm. longus, 1 mm.
latus; lobi laterales divergentes, lineari-oblongi, apice
subacuti, 1.25 mm. longi, 0.6 mm. lati; calcar valde
circinatim incurvatum, ex ore angusto cylindricum, apice
clavato-inflatum, obtusum, circiter 1 cm. longum. Co-
lumna \eviter incurvata, semiteres, fere 2 mm. longa,
apice truncata; androclinium leviter excavatum ; anthera
hemisphaerica, antice in appendicem obtusam producta;
pollinia ellipsoidea, 0.6 mm. longa, stipitibus linearibus
superne subspathulato-dilatatis 0.8 mm. longis, viscidiis
distinctis oblongis convexis 0.8 mm. longis; rostellum
productum, obtusum, viscidiis amotis trifidum; fovea
stigmatica orbicularis; ovarium circiter 2.5 mm. longum,
leviter 6-sulcatum.
Ucanva: Bunyoro, Budongo Forest, Siba block, by stream, June
1944, flowered Busingiro, July 1944, Eggeling 5386 (Typw).
This species is easily recognisable on account of the
very large leaves. In floral structure it resembles A.
gracillima (Rolfe) Summerh., but has a much shorter
clavate spur. An interesting feature is the widening of
the claw of the lip just in front of the spur opening to
produce two rounded angles which may be looked upon
as the ends of the decurrent sides of the column, though
there is no true foot in this genus.
5. Angraecopsis elliptica Swmmerhayes sp. nov.
affinis A. trifurcae (Rehb.f.) Schltr., sed foliis anguste
[ 250 |]
vel oblongo-ellipticis apice minus inaequilobatis, inflores-
centils brevioribus, floribus minoribus, labelli lobis later-
alibus quam intermedio satis minoribus, calcari breviore
differt.
Herba epiphytica, radicibus exceptis glaberrima. Caulis
brevis, usque ad 3 cm. longus, inferne radices flexuosas
ramosas pubescentes 2-3 mm. diametro emittens, superne
eirciter 4-foliatus. Molia disticha, recurvatim patentia;
vagina compressa, siccitate multistriata, 1-1.5 em. longa;
lamina anguste vel elongato-elliptica, apice paulo inae-
qualiter bilobulata, lobulis rotundatis vel obtusis longiore
vix 5 mm. longo, 4.5-10.5 cm. longa, 1.7—2.5 cm. lata,
supra secus medium canaliculata, subtus leviter carinata,
subearnosa. Inflorescentiae ex axillis foliorum delapsorum
exortae, simpliciter racemosae, 10-15 cm. longae, sub-
dense usque ad 18-florae; pedunculus gracilis, teres, 8-
10 cm. longus, vaginis paucis arctis acutis 3-4 mm. longis
instructus; rhachis levissime fractiflexa; bracteae ochre-
atae, apice acutae, usque ad 2 mm. longae. Fores in-
curvatim adscendentes, virides, hyalini; pedicellus cum
ovario circiter 5 mm. longus. Sepalwm intermedium
erectum vel leviter incurvatum, lanceolato-ovatum, acu-
tum, circiter 2.25 mm. longum et 1.25 mm. latum, tri-
nervium; sepala lateralia parallele porrecta, oblique et
incurvatim oblanceolata, circiter 4.75 mm. longa et 1.5
mm. lata, subquadrinervia. Petala triangularia, margine
antico rotundato-dilatata sepalo laterali adnata, apice
acuminata reflexa, circiter 2.25 mm. longa, basi 2 mm.
lata, trinervia. Labellum + incurvatum, ex ungue ob-
longo trilobatum, 4.5 mm. longum; lobus intermedius
lineari-lanceolatus, acutus, circiter 2.75 mm. longus, basi
0.75 mm. latus; lobi laterales lineares, divergentes, 1.6
mm. longi, basi 0.25 mm. lati; calear cylindricum, in-
curvatum, apice haud inflatum, circiter 5 mm. longum.
Columna brevis, crassa, circiter 1.25 mm. longa, apice
[ 251 ]
truncata; androclinium leviter excavatum; anthera non
visa; pollinia ellipsoideo-globosa, 0.5 mm. longa, stipiti-
bus linearibus 0.3 mm. longis, viscidiis distinctis quad-
ratis minutis; rostellum breviter productum, viscidiis
amotis breviter bidentatum ; ovarium leviter 6-suleatum,
circiter 2.5 mm. longum.
British Cameroons: Buea, 900 m. alt., 1947, Gregory 194 (Tyrer).
Chiefly recognisable by the relatively broad oblong-
elliptical leaves. In floral structure it is nearly related to
the Comoro Islands A. trifurca (Rehb.f.) Schltr. It is
a larger plant than either of the other three species from
this general region (A. ischnopus (Schltr.) Schltr., A.
dolabriformis (Rolfe) Schltr. and 4. tridens (Lindl. )
Schltr.) and may also be distinguished by the details of
the flowers. A specimen collected by J. W. Purseglove
(no. 2614) at Goma near Lake Kivu in the Belgian Con-
go, is probably also referable to this species, but the plants
are considerably smaller and unfortunately none of the
flowers is preserved completely.
6. Angraecopsis trifurca (Rchb.f) Schlechter in
Beih. Bot. Centralbl. 38, Abt. 2 (1915) 428 et in Beih.
Bot. Centralbl. 86, Abt. 2 (1918) 141 — ? Perrier de la
Bathie in Humbert, Fl. Madag. Orchid. 2 (1941) 82.
Aéranthus trifurcus Reichenbach filius in Flora 68
(1885) 540.
Mystacidium trifurcum Durand & Schinz, Consp. FI.
Afr. 5 (1895) 55.
Listrostachys trifurca Finet in Mém. Soc. Bot. France
9 (1907) 51, ? excl. deser. et ic.
Mystacidium Thouarsii Finet, |.c. 57, quoad descr. et
ic., excl. syn. et plantae Mauritianae et Borbonicae.
Angraecopsis Thouarsii H. Perrier in Humbert, FI.
Madag. Orchid. 2 (1941) 84, ewel. syn.
A. comorensis Summerhayes in Kew Bull. (1949) 443.
[ 252 ]
Comoro Istanps: Grande Comore, Humblot 450 (Tyre in Herb.
Vienna); Grande Comore, June 1899, Pobéguin.
Comparison of the type of Aéranthus trifurcus Rehb. f.
in the Vienna Herbarium (Humblot 450) with the ma-
terial collected by Pobéguin and described by Perrier
de la Bathie as Angraecopsis Thouarsii shows that they
are conspecific. The petals in the Reichenbach type are
trinerved, while the side lobes of the lip are nearly as
broad and not quite so long as the middle lobe, just as
in the Pobéguin specimens.
I am rather puzzled as to the true nature of the ma-
terial in the Paris Herbarium under the same number as
the type, Humblot 450, which was figured and described
by both Finet and Perrier. These drawings and descrip-
tions clearly do not agree with the type specimen at
Vienna, and it is therefore possible that Humblot col-
lected a mixture of species under his number 450. An
alternative explanation is that both French authors have
placed too much dependence on the drawings of the artist
C. Kastner which appear in both Finet’s plates and as
parts of Perrier de la Bathie’s illustrations. Examination
of drawings by this artist on sheets of other species in
the Paris Herbarium shows that they are sometimes
highly imaginative and do not agree with my dissections
of flowers from the same plants. It seems therefore pos-
sible that the drawings on the sheet of Humblot 450 are
also inaccurate and have led to a misinterpretation of the
material; clearly a re-examination of this specimen is
desirable.
7. Angraecopsis dolabriformis (2ol/fe) Schlechter
in Beih. Bot. Centralbl. 36, Abt. 2 (1918) 140 — Dandy
in Excell, Cat. Vase. Pl. S. Tomé (1944) 335.
Mystacidium dolabriforme Rolfe in Bolet. Soc. Broter.
9 (1891) 141 et in Dyer, FI. Trop. Afr. 7 (1897) 175
[ 2538 ]
— Durand & Schinz, Consp. Fl. Afr. 5 (1895) 52 —
Henriques in Bolet. Soc. Broter. 27 (1917) 183.
Angraecum dolabriforme Engler in Engler & Drude
Veget. der Erde 9, bd. 2 (1908) 419.
Sao Tome: no locality, Henriques, cult. in Hort. Coimbra in 1889
and in Herb. Kew (Typr).
This and the next species (4. tridens (Lindl.) Schitr.)
are clearly closely allied and further material may show
that they are conspecific. 4. dolabriformis has broader
leaves, more flowers in the inflorescence, slightly larger
flowers and a longer spur than 4. tridens, but it remains
to be seen if these are valid distinctions or merely rep-
resent plants of different degrees of robustness.
8. Angraecopsis tridens (Lind/.) Schlechter in
Beih. Bot. Centralbl. 36, Abt. 2 (1918) 141 — Summer-
hayes in Hutchinson and Dalziel, Fl. West Trop. Afr.
2 (1936) 462.
Angraecum tridens Lindley in Journ. Linn. Soc.
Lond. Bot. 6 (1862) 136.
Listrostachys tridens Reichenbach filius in Flora 48
(1865) 190— Durand & Schinz, Consp. Fl. Afr. 5
(1895) 50.
Epidorchis tridens O. Kuntze, Rev. Gen. Pl. (1891)
660.
Saccolabium occidentale Kraenzlin in Engl. Bot. Jahrb.
17 (1898) 57.
Angraecum occidentale Rolfe in Dyer, Fl. Trop. Afr.
7 (1897) 142.
Mystacidium tridens Rolfe, |.c. 174.
Angraecopsis occidentalis Schlechter in Beih. Bot. Cen-
tralbl. 36, Abt. 2 (1918) 140.
British Cameroons: Cameroon Mt., W. of Buea, 1240 m. alt., in
primeval forest, Sept. 21st 1891, Preuss 965.
Frernanvo Po: no locality, 1200 m. alt., Dec. 1860, Mann 646
(Tyrer).
[ 254 ]
The distinctions between this and A. dolabriformis
(Rolfe) Schltr. are mentioned under that species. Both
species are characterised by the short inflated spur.
9. Angraecopsis parviflora (Thou.) Schlechter,
Die Orchideen (1914) 601 et in Beih. Bot. Centralbl. 33,
Abt. 2 (1915) 428 et l.c. 86, Abt. 2 (1918) 140 — Perrier
de la Bathie in Humbert, Fl. Madag. Orchid. 2 (1941) 85.
Angraecum parviflorum Thouars, Orch. Iles Austr.
Afr. (1822) t. 60 — A. Richard, Orch. Iles France &
Bourbon (1828) 70— Bojer, Hort. Maurit. (1837) 317.
Aérobion parviflorum Sprengel, Syst. Veg. 8 (1826)
718.
Occeoclades parviflora Lindley, Gen. & Sp. Orch. PI.
(1838) 236.
Listrostachys parviflora 8S. Moore in Baker, F]. Maurit.
Seych. (1877) 855— Durand & Schinz, Consp. FI.
Afr. 5 (1895) 49.
Epidorchis parviflora O. Kuntze, Rev. Gen. Pl. (1891)
660.
Saccolabium parviflorum Cordemoy, Fl. [le Réunion
(1895) 197.
Mystacidium pedunculatum Rolfe in Dyer, FI. Trop.
Afr. 7 (1897) 175.
TANGANYIKA Terrirory: Kyimbila District, N. of Lake Nyasa,
Stolz K. 51.
Nyasatanp: Mt. Milanji, 1200 m. alt., McClounie 29; Manganja
Mts., Mt. Chiradzulu, Sept. 1861, Meller.
Portuguese East Arrica: Makua Country, Namuli Mts., Last.
Sournern Ruopesta: Umtali District, Vumba, Norseland, 1500 m.
alt., in kloof forest, March 1949, Wild 2799 (S. Rhodesian Gov. Herb.
No. 22724); Melsetter District, Lucite Valley, 1200 m. alt., Feb.
24th 1907, Johnson 175.
Mapaacascar: Manongarivo Mts., 1000 m. alt., in forest, March
1909, Perrier 1947.
Mauritius: no locality, Thouars 30 (Tyre); Grey; Bojer; Commer-
son (according to Bojer the species occurred in forests on the Pouce
and Picter. Both mountains and at Plaines-Wilhems).
[ 255 ]
Reunton: Mont St. Francois, S. of St. Denis, Richard 507; no lo-
eality, Richard 406; Boivin 1060; Herb. Drake (common at St. Benoit
according to Cordemoy).
Careful examination and comparison of all these speci-
mens indicate that they are referable to the same species.
The specimens from the African mainland are on the
whole larger than those from Mauritius and Réunion,
but there is practically no difference between the smallest
mainland plants and the largest island specimens. In
floral structure there is very close agreement, at any rate
as regards the two Southern Rhodesian gatherings of
which I have seen well preserved flowers.
Finet’ provides a description and figures of two gath-
erings (Sacleuw 764 and 765) from Portuguese East
Africa which he identifies with Rolfe’s Mystacidium pe-
dunculatum. Although there are no properly preserved
flowers on Rolfe’s material, in several cases the spur still
remains. This, even allowing for shrinkage, could not
possibly have exceeded 1 em. in length and was probably
nearer 7 mm., which is about the length of the spur in
Angraecopsis parviflora. On the other hand the flowers
figured by Finet have a spur which appears, from the
magnifications given, to be about 2 em. long with quite
a wide mouth. This certainly does not agree with the
shrivelled-up spurs on the capsules of MM. pedunculatum.
It therefore seems almost certain that Sacleux’s plants
are referable to a different species, which is correctly
placed in Mystacidium, with which they agree in floral
structure, especially the lip and rostellum-lobes.
It should be further pointed out that in no true species
of Mystacidium are the flowers bunched at the end of
the inflorescence with a long peduncle below, but are
spaced out evenly along the greater part of the inflores-
cence so that the peduncle is comparatively short. In
*Mém. Soc, Bot. France 9 (1907) 57, t. XII, fig. 1-12.
[ 256 ]
Angraecopsis, on the other hand, a long slender peduncle
is a very common occurrence, the flowers being closely
placed along the comparatively short rhachis. All this
evidence supports the view that the original Mystacidiwm
pedunculatum of Rolfe is correctly referred to Angrae-
copsis parviflora and has nothing to do with the plants
figured under the same name by Finet.
A. parviflora is clearly related to A. trifurca and A.
eliptica as regards the floral structure; the differences
between the species are indicated in the key.
10. Angraecopsis Pobeguinii (Finet) Perrier de
la Bathie in Humbert, Fl. Madag. Orchid. 2 (1941) 81,
fig. LI, 1-7.
Rhaphidorhynchus Pobeguinu Finet in Mém. Soe.
Bot. France 9 (1907) 41, t. VII, fig. 46-54.
Chamaeangis Pobeguiniu Schlechter in Beih. Bot. Cen-
tralbl. 38, Abt. 2 (1915) 426 et lc. 86, Abt. 2 (1918)
Lit,
Comoro Istanps: Grande Comore, June 14th 1899, Pobéguin (Tyre
in Herb. Paris).
This species is easily recognisable by the small flowers
and the very short side lobes to the lip. These stand out
almost at right angles so that the whole lip appears cross-
shaped; the apex of the middle lobe is much thickened
and fleshy.
In the original description and illustrations of the spe-
cies the two pollinia are shown as possessing a common
stipes and viscidium. I was not able to discover the pol-
linarium on examining a flower on the type specimen,
but I very much doubt the accuracy of Finet’s account
in this respect. One wonders if the imagination of the
artist, C. Kastner, is again here in evidence, as suggested
under A. trifurca. In this connection it may possibly be
significant that Perrier de la Bathie does not reproduce
[ 257 ]
the drawings of the pollinarium, though all the other
dissections from the type sheet appear on his plate;
neither does he mention the common stipes and viscidium
in his specific or generic descriptions. Instead he points
out that the two stipites and viscidia are sometimes coa-
lescent; I can confirm that this happens in dried mate-
rial of many species of the genus, though the viscidia are
clearly distinct in fresh or alcohol material.
Section 2. Cardiochilus Summerhayes
This section contains only one species, a small insig-
nificant plant with narrow leaves and few-flowered lax
inflorescences of small greenish-white flowers. In this
section the petals are only slightly differentiated from
the dorsal sepal and the line of attachment to the lateral
sepal is very short. The entire broadly ovate lip with
slightly cordate base is also a characteristic feature.
Nevertheless, the combination of characters present
seems to link the plant most closely with Angraecopsis
of which, however, it is clearly an aberrant species.
11. Angraecopsis pusilla Summerhayes sp. nov.
Herba epiphytica nana, tota usque ad 7 cm. alta.
Caulis erectus, brevis, usque ad 2 cm. longus, inferne
radices flexuosas carnosas glabras emittens, superne
crebre 5-9-foliatus. Folia + erecta vel adscendentia;
vagina compressa, 5-8 mm. longa, siccitate pluricostata;
lamina linearis vel lineari-ligulata, apice inaequaliter bi-
lobulata lobulo breviore fere nullo longiore rotundato,
1.5—5 cm. longa, 1.5-5 mm. lata,sectione leviter V-formis,
carnosula. Inflorescentiae ex axillis foliorum exortae,
graciles, superne laxe 4~—7-florae; pedunculus gracilis,
teres, 1-3 cm. longus, vaginis perpaucis arctis acutis
usque ad 4 mm. longis instructus; rhachis gracilis, fracti-
flexa; bracteae lanceolatae, dimidio inferiore vaginantes,
apice acutae, 1.5-2.5 mm. longae, extra carinatae. Flores
[ 258 ]
5-10 mm. distantes, erecto-patentes, albi, viridi-tincti;
pedicellus cum ovario 5-8 mm. longus, gracilis. Sepalum
intermedium leviter incurvatum, convexum, orbiculari-
ovatum, apice obtusissimum, circiter 1.75 mm. longum
et paulo angustius quam longius, subtrinervium; sepala
lateralia subspathulatim oblanceolata, curvata, apice ob-
tusa, circiter 2.75 mm. longa et 1 mm. lata, uninervia.
Petala \ate oblique elliptico-ovata, apice rotundata, mar-
gine antico paulo dilatata sepalo laterali adnata, circiter
2mm. longa et 1.6 mm. lata, binervia, nervis ramosis.
Labellum concavum, late ovatum, apice subacutum, basi
truncatum vel leviter cordatum, 2.5 mm. longum, ex-
planatum circiter 2 mm. latum, trinervium; calcar fere
rectum, ex ore angusto cylindricum, sursum sensim an-
gustatum, apice rotundatum, circiter 2.75 mm. longum.
Columna fere recta, brevis, crassa, subteres, 1 mm. longa;
androclinium leviter excavatum; anthera hemisphaerica,
antice brevissime producta; pollinia sphaeroidea, circiter
0.8 mm. diametro, stipitibus duobus linearibus polliniis
brevioribus, viscidiis duobus distinctis ellipticis parvis;
rostellum incurvatim porrectum, viscidiis amotis indivi-
sum; ovarium plurisulcatum, circiter 1.75 mm. longum.
Betaran Conco: Pare National Albert, between Nyamlagira and
Tsambene, 1700 m. alt., in forest, Jan. 16th 1945, Germain 3432
(Type in Herb. Brussels).
Described from both dried and alcohol material. Pos-
sibly with a wider distribution in the upper forests of the
Central African mountain region, but easily overlooked
on account of its small size and insignificant flowers.
Section 8. Coenadenium Summerhayes
The species of this section are characterised by the
possession of a common viscidium to the two pollinia,
though the stipites are distinct. Apart from the polli-
narium, the species exhibit the characteristic Angrae-
[ 259 ]
copsis floral facies more clearly than the species in the
last section, though not so markedly as the species of
sect. Mu-angraecopsis. They are all small plants with
rather short inflorescences in which the rhachis occupies
the upper half or more, the peduncle being comparatively
short. All the species are natives of East Tropical Africa.
12. Angraecopsis holochila Summerhayes in Bot.
Mus. Leafl. Harvard Univ. 12 (1945) 115.
Ucanva: Karamoja District, Napak, 2250 m. alt., in ravine forest,
May 28th 1940, Thomas 3646 (Tyre); same locality, 2190-2250 m.
alt., on trees and rocks, June 1950, Eggeling 5971.
Characterised by the almost entire lip and relatively
long spur. The large ovate very convex viscidium is
truneate and slightly emarginate in front; the stipites
arise laterally, are almost straight or slightly curved and
in length much exceed the diameter of the pollinia.
Further material in alcohol collected recently by Dr.
W. J. Eggeling has confirmed in all respects the striking
features of this delightful little species.
13. Angraecopsis amaniensis Swmmerhayes in
Bot. Mus. Leafl. Harvard Univ. 11 (1945) 259.
A. tenuicalear Summerhayes, l.c. 258.
Kenya Cotony: Mt. Elgon, south side, 2160 m, alt., in Podocarpus
forest, April 1943, coll. E. W. Carroll, comm. E. M. Tweedie 603.
Tancanyika Trrrtrory: West Usambara Mts., Shume, edge of
forest, March 25th 1942, Moreau 325; East Usambara Mts., Amani,
900 m. alt., on exotic conifers, July 4th 1941, Moreau 68 (Tyrr).
Since describing the above two species, I have seen
further material, including flowers in aleohol of Moreau
325, This additional material shows that, firstly, the sup-
posed difference in the thickness of the spurs of the two
species 1s non-existent and, secondly, the relative lengths
of the lateral and intermediate sepals are very variable,
intermediates occurring between the conditions described
[ 260 |
in my original accounts. It therefore seems more logical
to look upon the various gatherings as forms of one spe-
cies, the general floral structure and especially the column
showing very close agreement.
14. Angraecopsis breviloba Summerhayes in Bot.
Mus. Leafl. Harvard Univ. 11 (1945) 256.
Kenya Cotony: Mgong, 1800 m. alt., in forest, May 7th 1934,
C. Van Someran(Coryndon Mus. No. 3653); same locality and altitude,
May 1932, Mainwaring 1858 (Coryndon Mus. No. 3104); Ngong and
Langata Forest, 1800 m. alt., in dense shade on Acokanthera Schimperi,
April-May 1950, Piers 2; North Nyeri District, Nyeri, Thego River,
March-April 1941, Copley 26; Tana River basin, steppes of the Thika-
thika, July 16th 1893, Gregory.
TANGANYIKA Territory: Mondul, W. of Arusha, 1800 m. alt., May
6th 1942, Moreau 304 (Tyrer); Mbulu, June 7th 1942, Moreau 3044;
Kilimanjaro, Marangu, 1350 m. alt., in riverain forest, Nov. 27th
1941, Moreau 107; Lolbene Mt., 50 miles S. of Moshi, in forest, coll.
F. M. Page-Jones, cult. and flowered Amani, April 10th 1944, Moreau
815.
This and the previous species have the short stipites
of the pollinia geniculately incurved so that the pollinia
lie close to one another. The viscidium is more or less
rectangular with the front and back rounded-truncate.
The present species may be distinguished from A. aman-
zensis by the shorter much inflated spur.
SPECIES REJICIENDA
Angraecopsis Boutoni (Rehb.f.) Perrier de la Bathie
in Humbert, Fl. Madag. Orchid. 2 (1941) 82.
Angraecum Boutoni Reichenbach filius, Otia Bot.
Hamburg. (1881) 117.
Examination of the type specimen of this species
shows that it is not an Angraecopsis at all, but a Cham-
aeangis closely allied to, if not identical with, C. Harv-
otiana (Kraenzl.) Schltr. I am refraining, however, from
making a new combination in Chamaeangis, as | am not
certain of the correct specific epithet for the species
concerned.
[ 261 ]
ene SE, cate ee ee eos Geen ii ns te Gee seen Se eel ———— ee © _
BOTANICAL MUSEUM LEAFLETS
HARVARD UNIVERSITY
VoL. 14, No. 10
WHENCE CAME MAIZE TO ASIA?
BY
Paut C. MANGELSDORF AND Doucuas L. OLIvER’
THE intriguing question of pre-Columbian trans- Pacific
diffusion is with us again and modern Kon-Tikis now
compete with ancient Alexandrian fleets in the alleged
South Pacific Regatta. To the interested but confused
onlooker, it might appear that the specialists are divided
into two opposing camps with adventurous diffusionists
in bitter conflict with obstinate and reactionary propo-
nents of independent invention. To some of the individ-
ual specialists involved, the issue may, indeed, have this
emotional coloring. Basically, however, the lines are
drawn between those who are short on facts and use them
uncritically (although sometimes with superb imagina-
tion) and those who demand evidence and valid reason-
ing. In the paper on maize in Assam, which is the prin-
cipal basis for this critique, neither the authors’ selection
of facts nor their reasoning from those facts can, in our
opinion, support their theory of a pre-Columbian diffu-
sion of maize across the Pacific. In their favor, however,
it must be added that they do not profess to know in
which direction the diffusion took place.
The question as to which part of the world gave rise
to maize is by no means new, since it is one upon which
‘Associate Professor of Anthropology and Assistant Curator of
Oceanic Ethnology, Peabody Museum, Harvard University.
[ 263 ]
students of plants have differed for more than four cen-
turies. Sturtevant, a careful student of maize and of the
literature pertaining to it, compiled (1879) lists of names
of prominent herbalists and early botanists who had ex-
pressed opinions on the geographical origin of maize.
Among those who regarded maize as a plant of Old
World origin were: Bock, Ruellius, Fuchs, Sismondi,
Michaud, Gregory, Lonicer, Amoreux, Regnier, Viterbo,
Donicer, Tabernaemontanus, Bonafous, St. John, de
Turre, Daru, de Herbelot and Klippart. Equally im-
pressive is the roster of those who believed maize to be
an American plant: Dodoens, Camerarius, Matthioli,
Gerard, Ray, Parmentier, Descourtilz, de Candolle,
Humboldt, Darwin, F. Unger, Von Heer, de Jonnes,
Targioni-Tozzetti, Hooker, Figuer, Nuttall, Mrs. Somer-
ville and Flint. De Candolle’s case (1855) for the Ameri-
can origin of maize was so convincing and the evidence
which he marshalled to support his conclusions so substan-
tial that the possibility of an Old World origin of maize
has received little consideration from serious students in
more recent times. Especially has this been true since
Ascherson (1875) demonstrated the close relationship of
maize and teosinte, a plant unmistakably American.
The question of a pre-Columbian distribution of maize
in Asia has, however, been raised at least twice in this
century, first in 1909 by Collins, and now by Stonor and
Anderson (1949). The last named paper, since it pur-
ports to present new evidence on the question and coin-
ciding as it does with a fashionable new preoccupation
with the old problem of trans-Pacific diffusion of pre-
Columbian cultures, has been of particular interest. We
have been requested by a number of anthropologists to
review it and to evaluate the evidence on which it is
based. The paper has already been critically discussed
by Merrill (1950), who has questioned its principal con-
[ 264 |
clusions on general botanical and ethnological grounds,
and by Weatherwax (1950), who has quite appropriately
emphasized the important and fundamental differences
which exist between maize and its Asiatic relatives, and
has simultaneously emphasized the similarities among
the American Maydeae. We propose here to examine
critically the botanical and ethnographic evidence con-
cerned with maize upon which the far-reaching conclu-
sions of the authors rest.
Stonor and Anderson found the hill peoples of Assam
growing a group of maize varieties with characters said
to be ‘‘unusual’’ and utilizing them for food, feed, and
brewing. This maize which the authors designate as
‘*Race A”’ is said to be unknown in the coastal regions
of Asia, but rather widely distributed in Central Asia;
furthermore, it seems to resemble certain South Ameri-
can maize also designated as ‘‘Race A’’ which is common
archaeologically and certain features of which are still to
be found, although rarely, among living South American
varieties. These peculiar Asiatic varieties differ pro-
foundly from those of ‘‘Race C’’ which also occur in
both Asia and America, but which in Asia are largely
confined to the coastal regions. The introduction of Race
C to Asia is admittedly post-Columbian.
These facts are regarded by the authors as ‘‘fantastic, ”’
and it is stated that ‘‘any satisfying hypothesis must
border on the miraculous.’’ They conclude that maize
presumably ‘‘must either have originated in Asia or have
been taken there in pre-Columbian times. ’’
The evidence upon which these sweeping conclusions
rests falls into three categories: (1) botanical evidence
concerning the maize in question; (2) ethnographic evi-
dence on the maize-using tribes and the uses to which
maize is put; (3) supporting evidence from Polynesia in
favor of trans-Pacific diffusion. We shall consider only
[ 265 ]
the first two categories, since the third has already been
discussed by Merrill and will undoubtedly receive addi-
tional attention from others who are better qualified than
we to evaluate it.
The Botanical Evidence
The botanical evidence may in turn also be considered
under three categories: (a) that the Assamese maize is
unique and is related only to archaeological maize in
America; (b) that the present distribution of Races A
and C can be explained only in terms of a pre-Columbian
diffusion of one of them; (c) that Assamese maize re-
sembles sorghum.
Five varieties of maize from Assam are described. The
following ‘‘unusual’’ characters are said to typify one or
more of these varieties.
1. Uniformly green leaves, culms, silks and anthers.
2. Slender pendent tassel branches.
Straw-yellow endosperm; dull bluish-red aleurone.
Small isodiametrical kernels.
5. Many short internodes; lack of vegetative vigor.
6. Upright twisted tassel branches; short silks.
7. Tassel partly enclosed in a spathe-like cluster of
leaves.
8. A distinct bloom on the leaves and culm.’
9. Waxy pollen and endosperm.
These characteristics, although not common, are never-
theless well-known to those familiar with the great di-
versity of maize in Latin America. Separately most of
them are widely distributed not only in South America,
but also in Central America and Mexico. Even in com-
? This character was not specifically included in their list, but is
mentioned separately as one of the characteristics of the variety Late
Sidewise.
[ 266 ]
bination they are by no means unique. In this connec-
tion the following observations made largely on varieties
of maize in the collections of the senior author are per-
tinent.
1. Uniform greenness resulting from a complete lack
of anthocyanin coloration is indeed almost unknown, at
least in pure form, in the indigenous maize varieties of
Latin America. Virtually all maize, however, has at least
a trace of anthocyanin color in the seedlings where it
occurs in the coleoptile or leaf sheath, in the tip of the
leaf blade, or along the leaf margins. Consequently, if
the Assamese varieties are actually completely lacking
in anthocyanin pigmentation, they are indeed almost
unique. Unfortunately, the authors do not report speci-
fically on anthocyanin color in the seedlings, stating only
that: ‘‘The group as a whole had a strong tendency to
green silks, green anthers, green leaves, and green culms. ”’
Plants of this general description are the product of two
recessive alleles at the B and P/ loci on chromosomes 2
and 6 respectively and of one of the lower alleles at the
R locus on chromosome 10. Such plants are not common,
but occur regularly throughout Latin America. In 1950,
we grew 513 collections of corn from sixteen Latin-
American countries. Among these were 27 varieties
which contained plants lacking in anthocyanin color in
the leaves, culms, silks and anthers. These occurred in
collections from Mexico, Guatemala, Honduras, Nica-
ragua, Colombia, Ecuador, Peru, Venezuela, Brazil,
Uruguay and Paraguay. They had their highest fre-
quency in the varieties of eastern South America. Of 40
varieties from Venezuela, Brazil, Paraguay and Uruguay
which were studied in 1950, eight, or one variety in five,
contained some uniformly green plants compared to one
variety in 19 for the group as a whole. So far as antho-
eyanin coloration alone is concerned, the Assamese vari-
[ 267 ]
eties, therefore, have their affinities in the corn varieties
of eastern South America.
In so far as uniform greenness has any bearing upon
the origin of maize, it points to Asia, not as a center of
origin, but as a peripheral region where recessive genes
have become ‘‘emancipated”’ through the process of
‘genetic drift.’’ The counterpart of this situation occurs
in rice which in the United States is represented largely
by uncolored (green) varieties, but which has colored
varieties in the Old World (Jones, 1980). In rice, as in
maize, at least three loci are involved in anthocyanin
coloration.
2. Pendent tassels, like lack of anthocyanin color, al-
though not common, are found in varieties from several
countries, including Mexico, Guatemala, Honduras,
Costa Rica, Nicaragua, Venezuela, Colombia, Peru and
Ecuador. The combination of pendent tassels and all-
green plants occurs in only seven of these nine countries,
not having been found in Costa Rica and Honduras.
All-green plants with strongly pendent tassels are most
common in Colombia where many other of the ‘‘unusual’”’
characters of the Assamese maize also occur,
3. Straw-colored endosperm is not at all unusual
among non-Tripsacoid varieties. Dull bluish-red aleurone
is the product of superimposing blue aleurone on waxy
endosperm and is a characteristic quite familiar to the
majority of practicing maize geneticists.
4. Small isodiametrical kernels are characteristic of
many South American pop corns. Dr. Anderson kindly
sent us kernels of several of the Assamese varieties. It
was possible to match more than half of these in size,
shape and color from a single collection of Colombian
pop corns.
5. Lack of vegetative vigor, manifested especially by
the slowness of the tassel to reach the pollen-shedding
[ 268 ]
stage after emergence has begun, is characteristic of many
South American and some Mexican varieties. It is surely
of little significance in the Assamese varieties which are
obviously rather highly inbred, if the statements of the
authors are correct regarding their uniformity and the
fact that they are sometimes grown as single plants
among other cereals. Lack of vigor is characteristic of
many American inbred strains.
We have no data on internode pattern.
6. We have no observations on upright tassel branches
and short silks.
7. The spathe-like cluster of leaves partly enclosing
the tassel is not uncommon in Colombian varieties.
8. The grayish bloom which shows some resemblance
to the bloom characteristic of sorghum (and many other
grasses) occurs in our collection only on varieties from
Colombia.
9. Perhaps the most important ‘‘unusual’’ character-
istic of the Assamese maize is the waxy endosperm which
occurs in several varieties. It was the discovery of this
character in Chinese maize which led Collins (1909) to
reopen the question of a pre-Columbian distribution of
maize in Asia.
Waxy endosperm is a simple Mendelian character in
maize which affects the chemical composition. The starch
of waxy maize is composed exclusively of amylopectin,
while that of non-waxy varieties contains both amylose
and amylopectin. Waxy varieties of maize are unknown
in pure form in America, but the waxy character itself
has been discovered in non-waxy varieties: in a New
England flint corn by Mangelsdorf (1924) and ina South
American variety by Breggar (1928). Bear (1944) has
found that waxy endosperm is not an uncommon mutant
in Corn-Belt dent corn varieties. He found three sepa-
rate mutations in three consecutive years in a total pop-
[ 269 ]
EXPLANATION OF THE ILLUSTRATION
Prare XLVIII. A pendent tassel of one of the
Colombian pop-corn varieties which resembles in
several characteristics the Assamese race Late Side-
wise described by Stonor and Anderson. Note, at
lower left, the sterile spikelets resulting from the
failure of the tassel to emerge completely from the
spathe-like sheath. Note, at right, the lax cen-
tral spike with spikelets borne in whorls at widely
separated nodes. Note the solitary spikelets at the
ends of several branches.
PLiateE XLVIII
ulation of some 100,000 selfed ears. It is of interest to
note that in the case of one of these mutations Bear
found one ear pure for waxy in the line in which waxy
endosperm was first noticed. This pure waxy ear was
the product of Mendelian segregation and not of human
selection.
In Asia, waxy maize is widespread. Collins (1909, 1920)
has reported its occurrence in China, Burma and the
Philippines. Kuleshov (1928) states that it is spread from
5° to 45° north latitude in Asia.
Is there any significance in the fact that a gene which
is comparatively rare in American maize should be wide-
spread in Asia’ Certainly there is none from the stand-
point of the time required for waxy varieties to become
established. It is not at all uncommon for recessive genes
which are rare at the center of a plant’s origin to become
common somewhere at the periphery of its spread. This
is anatural consequence of the process already mentioned,
‘‘oenetic drift,’ in which recessives with a low frequency
may rapidly attain a high frequency as the result of
sampling and without the intervention of either natural
or artificial selection. So far as waxy endosperm has any
bearing upon the origin of maize, it, like the all-green
plants discussed above, points to Asia as a peripheral
region rather than as a center of origin.
There is, however, undoubtedly other significance in
the fact that waxy maize occurs so commonly in a part
of the world which also possesses waxy varieties of rice,
sorghum and millet. The obvious explanation is that the
people of Asia, being familiar with waxy (glutinous)
varieties of other cereals and accustomed to using them
for special purposes, recognized the waxy character in
maize, when that cereal was introduced, and purposely
isolated varieties pure for the waxy condition’. Because
* Burkill (1935) questions, however, whether waxy maize was ever
[ 271 ]
waxy endosperm is arecessive, this task would have been
simple and well within the abilities of even the most
primitive hill peoples. Indeed, the practice of growing
maize as single plants among other cereals, reported by
Stonor and Anderson, would promote self-pollination,
and in any stock in which the waxy gene occurred would
inevitably lead in a very short time to the establishment
of pure waxy varieties whose special properties people
accustomed to the waxy character in other cereals could
hardly fail to recognize. Man’s part in the establishment
of waxy varieties of maize in Asia need have been no
greater than a recognition of this type and a willingness
to preserve it once it was presented to him as the product
of random sampling.
But much more important than the individual charac-
ters of the Assamese maize is the fact that these characters
occur as a “‘complex’’ in Asia: a complex which is said
to be rare in South America and ‘‘nothing like it’’ to be
known in Mexico, Guatemala or other parts of Central
America. How accurate is this statement and how valid
the conclusions regarding the uniqueness of Assamese
maize /
The complex of characters in its entirety has not been
reported from Mexico and Central America, but does
(if we exclude waxy endosperm as an integral part of the
complex) occur in South America. The authors them-
selves mention two varieties from Chile, one from Argen-
tina, and several from Bolivia which have a number of
features in common with the Assamese maize, and they
quote Brieger as noting these characteristics in other
parts of South America. The most unusual of the As-
samese varieties, called Late Sidewise, which is said to look
selected for its waxiness and suggests that it was preserved only be-
cause of the peculiar suitability of its sheaths for cheroots. This prob-
lem merits further study.
[ 272 |
‘unlike anything previously reported for Zea Mays’’ has
strong affinities, if not exact counterparts, among the
living varieties of Colombia. In 1949, we noted, in a
group of pop-corn varieties sent from the Department
of Caldas in Colombia by Dr. J. G. Hawkes, in a pop
corn received from Dr. R. E. Schultes collected slightly
north of Buenaventura, and in pop corn purchased by
the senior author in the market in Bogota, practically
all of the characteristics mentioned in the description of
Late Sidewise, including the bluish-green color of the
leaves and the culms and the distinct bloom which lends
to the plants a superficial resemblance to sorghum.
These Colombian pop corns are of unusual ethno-
graphic and botanical interest. They are known locally
as maiz indio and, according to Dr. Hawkes, they are
grown by the Indians in a primitive way, the seed being
sown broadeast and the crop receiving no weeding or
cultivation from the time of planting until harvest. A
similar statement about the method of sowing accom-
panied the collection made by Dr. Schultes. One of the
ears (No. 13855) of the Colombian pop corn (inadvertently
shelled off and put into cold storage before a photograph
could be made) was almost a duplicate of the ear (Stonor
No. 18) illustrated in Plate 21 of Stonor and Anderson.
Of seven distinct samples of kernels of Assamese maize
sent to the senior author by Dr. Anderson, five could
be matched almost exactly in size, shape and color with
Colombian pop corns. The pendent tassels of these corns
are illustrated in Plate XLVIII. They are of further bo-
tanical interest because of the slender, lax central spikes
of the tassel, on which the spikelets are borne in distinct
whorls separated by conspicuous internodes. The tips of
the tassel branches often bear solitary staminate spike-
lets. The plants tiller profusely and have numerous elon-
gated lateral branches. It was noted in 1949 that: ‘*these
[ 273 ]
plants look like corn-teosinte hybrids,*’ but there is no
other indication that they are the product of recent teo-
sinte contamination.
The complex of characters described for the Late
Sidewise maize of Assam is, as Stonor and Anderson con-
cluded, an unusual one. It is, however, not unique nor
confined to Asia. The fact that it occurs in Asia is not,
in our opinion, evidence either that maize originated
there or that it was taken there in pre-Columbian times.
The maize of Italy is in some respects as unusual as the
maize of Assam, but it, too, has American affinities. No
maize has yet been found in any part of the Old World
which does not have its counterparts in America. The
maize of Assam is no exception to this general rule.
The authors make much of the fact that the predom-
inating maize of Assam, Race A, is not represented in
collections of the maize from the Asiatic coast, and they
ask how such a race of maize could have gotten to a
number of isolated hill areas in Asia without leaving a
very definite record along the coast. ‘‘That maize,’ they
state, ‘‘could in post-Columbian times have spread to
each of these various hinterlands without entering into
the economies of the more civilized people who would
have handed it on almost passes belief.’’ And again,
‘To believe that in post-Columbian times maize could
have penetrated not only to the Naga but to the hill
tribes of Upper Burma, and of Siam, to the Lolo in cen-
tral Asia, to the aborigines of Hainan, to the hill peoples
of Sikkim, and to the interior of New Guinea, in each
case passing over the more civilized peoples along the
coast is beyond credulity.’
For us it is more difficult to believe that maize could
have occurred in pre-Columbian times in all of these
places, as well asin the coastal regions where it has now
presumably disappeared; and perhaps throughout Cen-
[ 274 |
tral Asia from Persia and Turkestan to Tibet and Siberia
where it now occurs, without leaving a single prehistoric
trace of any kind. Yet there is no tangible evidence of
the existence of maize in Asia or any other part of the Old
World before 1492. When we consider how thoroughly
other economic plants were treated in the extensive
ancient literature of Asia and the Near East, and how
popular maize became as a cultivated plant and as a sub-
ject for artistic treatment after the discovery of America,
it taxes our credulity to believe that all of the civilized
people of the Old World could have remained ignorant
of a food plant at once so widely distributed, so peculiar
in its characteristics and so useful to mankind.
Burkill (1935), probably the leading authority on the
economic plants of the Far East, came to a similar con-
clusion. He states: ‘‘The strongest reason against the
belief [of a pre-Columbian distribution of maize in China]
lies in the unanswerable argument that no plant of such
value could have remained hidden in the Far East, if
there.’”’
Actually the absence of Race A in the coastal regions
of Asia is not difficult to explain, if indeed it requires
explanation. In the first place, its absence among the
very limited collections so far made from the coastal re-
gions of Asia is far from conclusive proof that it does not
occur. But if we assume for the purpose of discussion
that Race A actually is absent at low altitudes in Asia,
then there are several possible explanations which do not
require the assumption of pre-Columbian diffusion. An
obvious one is that the more productive Tripsacoid vari-
eties of the second race, Race C, have already, in coastal
regions, largely replaced the non-vigorous unproductive
varieties of Race A, earlier introduced, as they are per-
haps in the process of doing in the hills where Race C
now also occurs along with Race A. The counterpart of
[ 275 ]
this process can be observed in many parts of North and
South America today.
A second possible answer, probably the correct one,
was given, forty years before the question was raised, by
Laufer who, as the result of his scholarly historical stud-
ies, concluded that maize came into China, not from
across the Pacific, but overland through Tibet from India.
This conclusion is quite in harmony with the facts of both
history and geography. Colombia, for example, where
living counterparts of the Assamese maize are now known
to occur, is actually appreciably nearer to Assam via the
Caribbean Sea, the Atlantic Ocean and Africa than via
the Pacific. Furthermore, the first route, being more
largely a land route, does not demand the fabulous feats
of navigation on the part of pre-Columbian people which
the second does.
This does not mean that Laufer’s conclusions on the
introduction of maize into Asia are necessarily completely
correct and final. However, until new evidence in con-
flict with them is brought forward, they furnish a satis-
factory explanation of the facts now at our command.
This is recognized by Stonor and Anderson who state
that accepting the morphological similarity of American
and Asiatic maize as a premise, Laufer ‘‘could have come
only to the conclusion he finally reached: that maize
somehow got to Indian ports at an early post-Columbian
date and spread overland via various primitive peoples to
China.’ Since it can now be shown that the Assamese
maize is indeed similar to American maize, the evidence
presented by Stonor and Anderson tends to confirm
rather than to contradict Laufer’s conclusions.
In his part of their joint paper, Anderson emphasizes
the resemblance of the Assamese maize in several char-
acteristics to sorghum, the implication apparently being
that this resemblance has some bearing upon the possi-
[ 276 ]
bility of an Old World origin of maize, since sorghum
is undeniably an Old World cereal. Actually the alleged
resemblances of Assamese maize to sorghum are either
superficial or are examples of the well-known phenome-
non of parallel variation which is especially well exem-
plified among plants by the cultivated cereals and among
animals by the rodents.
In the category of superficial resemblances are the
isodiametric straw-colored or dull blue kernels. Maize
kernels when not crowded tend to be spherical, and the
fact that kernels of Assamese maize approach this gen-
eral shape merely indicates that they are borne on ears
on which the kernels are not crowded. This is true of
many varieties of South American maize. Furthermore,
any variety of maize will produce spherical sorghum-like
kernels when it bears kernels in the tassel, as practically
all varieties are capable of doing when grown in small
pots in the greenhouse or when otherwise stunted.
The resemblance in kernel color between the Assam-
ese maize and sorghum is meaningless, since entirely
different color-bearing tissues are involved in the two
plants. The yellow and blue colors of the Assamese
maize are endosperm and aleurone colors respectively
and occur in triploid tissue resulting from the process of
double fertilization which is characteristic of the Angio-
sperm seed. The colors of sorghum kernels occur in the
pericarp and nucellar layer both of which are diploid ma-
ternal tissues (Swanson, 1928). So far as we know, en-
dosperm and aleurone colors have never been reported
in sorghum.
The resemblances of Assamese maize to sorghum in
lacking anthocyanin pigmentation and in possessing a
distinct bloom are nothing more than typical examples
of parallel variations in cereals and other cultivated
grasses such as sugar cane, in which variations in antho-
[ 277 ]
cyanin coloration and the presence or absence of bloom
are the rule rather than the exception. To find the grayish
bloom, one need go no further than corn’s closest rela-
tive, teosinte.
Finally, to link the Assamese maize with the ancient
Bat-Cave corn described by Mangelsdorf and Smith
(1949) by comparing both to sorghum is scarcely justi-
fied. The Bat-Cave corn resembles sorghum in its small
kernels, long glumes, and the fact that the upper glumes
are as long or longer than the lower. These, however,
are characteristics found in many varieties of pod corn.
To emphasize the resemblance of maize to sorghum or
to any other of the Old World relatives of maize without
also calling attention to the existence of profound and
fundamental botanical differences is to present a mislead-
ing picture. Maize does, indeed, resemble sorghum in its
general growth habit as well as in chromosome number,
and it resembles its Asiatic relatives Coix, Schlerachne,
Chionachne and Polytoca in being monoecious. It differs
from all of these, however, either in the development of
its florets or in the nature of its fruit case. Weatherwax,
some years ago (1926), called attention to the superfici-
ality of some of the resemblances between maize and its
Oriental relatives. He states:
In all the Maydeae the fruit is wholly or partly covered by an
indurated shell, which is an especially attractive superficial indica-
tion of relationship. Its relative absence in Zea may be explained
by the unusually complicated covering of husks, or as a result of
conscious selection by man. But this general occurrence of a hard
shell is a deceptive analogy, rather than a homology. The indurated
structure is a combination of a glume and an alveolus of the rachis
in Tripsacum and Euchlaena [as well as in Zea], a spathe in Coix,
and a glume in Polytoca, Schlerachne and Chionachne. A tendency
toward induration of something connected with the fruit seems,
therefore, to be all that the genera have in common, and this is
possessed by so many other genera of grasses as to be of little sig-
nificance in determining tribal relationships.
[ 278 |
Geographically the Maydeae are sharply divided into two groups,
one in each hemisphere, and neither has ever made its way into
the field of the other without the help of man. On the other hand,
all the genera of each group overlap sufficiently in distribution to
suggest an American progenitor and another in Australasia.
Weatherwax might quite justifiably have emphasized
even more than he did the close resemblance, morpho-
logically, of maize to its two American relatives, teosinte
and ‘Tripsacum. ‘True, its close relationship to teosinte
may be of little significance if teosinte is, as has been
suggested (Mangelsdorf and Reeves, 1939), a hybrid of
maize and ‘T'ripsacum. But the resemblance of maize and
Tripsacum, an indigenous American species which is
widely distributed in both North and South America,
is greater than is commonly recognized and is certainly
highly significant. In both genera one floret in each pis-
tillate spikelet is suppressed and in both it is the lower
floret which undergoes such suppression. In both genera
the caryopsis is either enclosed, or surrounded at the
base, by a structure which is made up of a segment of
the rachis containing an alveolus, and the glumes. In
Tripsacum the glumes are indurated while in maize they
are often membranous or fleshy, but there is evidence
from maize-teosinte crosses that this difference is in some
cases a simple Mendelian one of the same general mag-
nitude as that which distinguishes sweet corn from field
corn. Maize normally bears paired pistillate spikelets and
Tripsacum solitary ones, but paired spikelets have been
observed in Tripsacum by Dr. Cutler and solitary spike-
lets in maize by Hepperly (1949), so that discontinuity
between the two plants in these characters is not com-
plete. Maize is an annual and Tripsacum a perennial
possessing several characters normally associated with the
perennial habit. The distinction is not of profound im-
portance since annual and perennial species are some-
times found within the same genus. The resemblance to
[ 279 ]
Tripsacum of homozygous pod corn in bearing staminate
spikelets above and pistillate spikelets below on the
branches of the tassel is especially impressive.
Finally, it is possible, in spite of differences in chromo-
some number, to hybridize maize and ‘Tripsacum and to
demonstrate interchange between their chromosomes.
There is abundant circumstantial evidence that such hy-
bridization has occurred in the past and that it has been
an important factor in the evolution of maize under
domestication.
The closeness of relationship between maize and its
American relatives seems to us to be far more important
than the fact that maize has a larger number of relatives
in Asia than in America.
In short, there is nothing in the botanical evidence of
Stonor and Anderson in the three categories considered
to invalidate the widely-held and well-supported opinion
that maize is an American plant and there is nothing
which indicates to us that maize was taken across the
Pacific to Asia before 1492.
The Ethnographic Evidence
The ethnographic evidence of Stonor and Anderson,
like the botanical evidence, comprises several distinct
categories: (a) evidence concerned with legends and tra-
ditions; (b) names applied to maize ; (c) the uses to which
maize is put; (d) the role of maize in the economy of
the people.
Stonor in his part of the joint paper gives unwarranted
credence, we think, to statements by natives that maize
is a very old crop in the region studied. For example:
‘The Angamis I have talked to simply state that they
have grown maize from time immemorial.”’ ‘“The Abor
tribes simply state that they have always had maize
among their crops.”’ How simplified ethnology would
{ 280 |
be if all native informants were indeed as historically
reliable as the Assamese are inferred to be. But what
a confusing picture of the origin of cultivated plants one
would gain by giving credence to such unsupported state-
ments. ‘This conviction on the part of native peoples that
they have ‘‘always”’ had a certain plant is by no means
confined to the Assamese. Dr. Carl Coons tells us that
the natives of Albania are convinced that they have al-
ways had tobacco. ‘The native peoples of the Near East
are quite certain that they have always had squashes; and
Irish peasants, if the question were put to them, would
no doubt answer that they have always had the potato.
The Indians of Central America are convinced that they
have always cultivated the banana, a fact which would
undoubtedly be regarded by some as evidence of early
trans-Pacific diffusion. But the same Indians, or their
neighbors at slightly higher altitudes, are equally sure
that they have always had the broad bean Vicia Faba,
one of Europe’s principal leguminous food plants.
Legends to account for the origin of rice are regarded
by Stonor as significant, since there is ‘‘no legend known
to account for the origin of the other cereals; millet,
maize and Job’s-tears, the inference being that rice is
more recent while the others are lost in the mists of an-
tiquity.’’ ‘This is, to say the least, an unusual criterion
of ethnological age.
Stonor found distinct names for maize in several of the
tribes surveyed and regarded this as ‘‘everywhere indica-
tive of a respectable age,’’ and he did not consider the
case weakened in instances where the tribal name indi-
cates that it was borrowed from a neighboring people,
since ‘‘the generalized name could be based on a variety
got from the tribe in question and which supplanted
older and more indigenous types.’” The fact that there
is no evidence of any kind of ‘‘older and more indige-
[ 281 ]
nous types’’ seems to be of no importance in answering
the question of the antiquity of maize in Assam. Its
antiquity seems to be a basic assumption to which the
author clings despite conflicting evidence.
Particular emphasis is placed upon the multiple uses
of maize among the hill tribes. These are: (1) a catch
crop eaten while the grain is soft; (2) stored for winter
food either as the main crop or as a reserve secondary to
rice; (8) for beer making; (4) for pop corn; (5) for pig
food; (6) as an article of trade outside the village.
Obviously the authors do not have a high opinion of the
capabilities of pre-literate peoples: ‘*‘’T'o have these con-
servative people somehow learning to use maize as a pop
corn and as a green corn and as a cereal for brewing, to
have them growing types of maize which are similar to
each other yet rare or unknown in the New World puts
the burden of proof on any one who would ascribe all this
development to separate post-Columbian acquisitions. ”’
How else would primitive people be expected to use
maize! If they use maize at all they must surely use it
for food and once used for food it would be likely to be
used both green and ripe, as it is in all other parts of the
world where maize is grown. And if the mature maize
is small and hard and capable of popping, how much in-
genuity is required to put grains of maize “‘into the glow-
ing embers of the fire,’ or ‘‘in the edge of the house
fire’’ picking them out with bamboo tongs as they burst ?
How often has the discovery been made independently
that small hard kernels of maize will pop when exposed
to heat? Is there any greater significance in the fact that
the Assamese use maize for popping than in the fact that
Asiatic people in general use seeds of species of Amaran-
thus for that purpose, or the fact that people throughout
Latin America use hard-seeded varieties of sorghum, an
African plant, for popping?
[ 282 |
And what is so strange about using maize for brewing?
Practically all of the cereals have been used for brewing
in practically all parts of the world where the art of brew-
ing has been practiced. It would be much stranger if the
Assamese, ‘“‘conservative’’ as they are said to be, did not
use maize for this purpose.
Or does the feeding of maize to pigs call for an expla-
nation? Given both maize and pigs, the problem quickly
becomes one which the pig itself is likely to solve with-
out much help from man. Domestic pigs have shared in
man’s principal carbohydrate foodstuffs since time im-
memorial and no great amount of ingenuity on the part
of man is needed to establish this relationship.
The use of maize as an article of trade is too obvious
to need comment.
A special use of maize in religious rituals is also re-
garded as a measure of antiquity. For example: ‘‘the
dance of the Lakhers, the use of maize in funeral rites
among the Lushais in deliberate preference to rice, its
importance as a votive offering among the Monbas, the
part it plays among the agricultural ritual of the Rengma
Nagas, and the existence of aspecial tutelary deity among
the Chang Nagas, all point to its being a well-established
crop, the more so since primitive peoples with animistic
religion are invariably shy of incorporating new crops
into their agricultural ritual.’’ Nor is negative evidence
allowed to weaken this case: ‘‘I have asked members of
the [Monba] tribe if they have any special rites, dances
or festivals for their maize, and in all instances this was
denied. I would not, however, like to state categorically
that my informants were accurate. In dealings with tribal
peoples knowledge of religious custom can only be got
by long and close acquaintance or direct observation. ”’
And, as the author himself admits, ‘‘ My notes on the
religious aspect are particularly scanty.’” How unfortu-
[ 283 ]
nate that the author does not display the same admirable
caution towards his other kinds of ethnographic data.
The position of maize relative to other crops in the
economy of the hill peoples does not seem to us to have
the significance which the authors attach to it. Maize,
like millet and Job’s-tears, is subordinated to rice among
peoples living at lower altitudes, but is said to be ‘‘of
more importance to the tribes living at high altitudes. ”’
While we can agree with the authors that this probably
is ‘a state of affairs not unconnected with absence of rice
varieties suitable for cold elevations, ’’ it is difficult to see
how this has any bearing on their case. In view of the
ethnological sequence in this region of Asia, it is not sur-
prising to learn that rice is a relatively recent introduction
among some of these tribes in spite of the fact that rice is
an ancient Asiatic food crop. We cannot, however, put
these circumstances together to conclude, as these au-
thors have done, that maize is necessarily pre-Columbian
in this region.
Laufer concluded some years ago that maize may have
reached China as early as 1540. Goodrich (1988) dates
the first Chinese reference to it at 1573. Some 400 years
have now elapsed since maize came to Asia. It surely
does not tax an anthropologist’s credulity to believe that
the Assamese and their neighbors, however conservative,
have within this period learned or rediscovered or adapted
to their own purposes several of the most obvious ways
of using maize. Wonder would have been aroused if
they had not.
Parallels between Maize in Asia and the Potato
in Ireland
To those who are astonished at the extent to which
maize is grownin Asia and the number of uses to which
it is put and who feel that more than four centuries must
[ 284 ]
be allowed for the plant to have established itself so firmly
in the economy of backward peoples, a study of the his-
tory of other cultivated plants may be revealing. Espe-
cially illuminating is a recent scholarly treatise by Sala-
man (1949) on the potato, and of particular interest are
those chapters concerned with its history in Ireland.
Introduced into Ireland between 1586 and 1588, the
potato had, within fifty years of its introduction, ‘‘be-
come the universal and staple article of the peoples’ food
in the greater part of the island.”’ Many indigenous
names were invented for it: pratie, tata, murphy, croker
and buntata. Many superstitions and social and religious
customs grew up in connection with its culture. The
potato was used not only for food in a variety of ways,
but also medicinally and in the preparation of an alco-
holic beverage. It was not only food for man, but also
provided nourishment for all of his domestic animals,
‘‘the pig taking his share as readily as the wife, the cocks,
hens, turkies, geese, the cur, the cat, and perhaps the
cow — and all partaking of the same dish.”’
‘*So completely had the potato woven itself into the
web of the life and thought of the people’’ that they
were immune to warnings of crop failures, and ‘‘no more
attention was given to such warnings than would have
been the case had they been told that the rains would
cease to fall from heaven.’’ The potato, like the sun and
the stars and the rain, had ‘‘always’’ been with them.
It may be argued that the potato became a part of
the Irish culture so rapidly only because the Irish were
already an advanced people. This apparently is not the
case. The state of agriculture in Ireland in the sixteenth
century was very primitive indeed and, in Salaman’s
opinion, it was this very backwardness of the Irish accom-
panied by general devastation and misery which lead to
the breakdown of prejudice against, and the rapid accep-
[ 285 ]
tance of, anew food. Certainly the potato became much
more quickly established in Ireland than in nearby Eng-
land where it had been introduced even earlier. Also, it
is known that both the Germans and the Scots strongly
resisted the potato until famine dispelled their prejudice.
Perhaps the rapidity with which a new plant is adopted
by any people is less a function of their progressiveness
than of their need. Seen from this viewpoint, the rapid
spread of maize in Asia is not at all astonishing. In the
light of the history of the potato in Ireland, post-Colum-
bian time has been ample, and more than ample, for the
introduction of maize into Asia and for its establishment
as a staple crop.
The Origin of New World Cultivated Cotton
and Its Bearing on Asiatic Maize
Stonor and Anderson, to support their argument for an
origin or a pre-Columbian distribution of maize in Asia,
cite the hypothesis of Hutchinson, Silow and Stephens
(1947) which postulates that the New World cultivated
cottons are tetraploid hybrids of a wild American diploid,
probably Gossypium Raimond, and a cultivated diploid,
G. arboreum, introduced from Asia by man crossing the
Pacific after the invention of agriculture in Asia. This
hypothesis has also been cited by others (Carter, 1950;
Zelinsky, 1950) as evidence of pre-Columbian trans-
Pacific diffusion. It should perhaps be pointed out that
many botanists, including the senior author of this paper,
although they recognize the hypothesis as stimulating
and provocative, are quite critical of it on genetic and
botanical grounds. The reasons for this are several.
First, there isno more need of explaining the distribu-
tion of the Old and New World cottons in terms of
man’s peregrinations than there is of accounting for the
range of numerous other genera which have a similar
[ 286 ]
geographic distribution. Indeed, if the differentiation of
cotton species is to be explained in terms of man’s move-
ments, then there are other genera which are not culti-
vated, in which speciation ought likewise to be so ex-
plained ; a procedure which would soon reduce the thesis
to an absurdity.
Secondly, their classification of Gossypium, based upon
the assumption of a recent origin of the New World tet-
raploids, is not in harmony with some of the sound tax-
onomic conclusions of earlier students. For example, the
endemic cotton of the Galapagos Islands, formerly re-
garded as a good species, G. Darwinii, is now treated as
a variety of the mainland cotton G. barbadense.
Finally, the endemic wild tetraploid cotton of Hawaii,
G. tomentosum, presumably derived from the American
tetraploid, presents an almost insuperable difficulty to
the entire hypothesis. How could the Hawaiian cotton,
in a few thousand years or less, have become so differ-
entiated from the mainland allotetraploids that it is now
generally regarded as a distinct species, since it differs in
many characteristics, and since there is a high incidence
of seedling mortality in the F, when G. tomentosum is
crossed with the American species G. hirsutum. The
genetic gap between the Hawaiian tetraploid and the
American tetraploids is perhaps a fourth to a half as
great as the gap between the American and Asiatic dip-
loids, yet the differentiation in the one case is supposed
to have required only a few thousand years, in the other,
since it is assumed to have begun in the Cretaceous, some
120 million years. Differentiation of species does not,
of course, proceed uniformly in time and space and the
degree of differentiation is not a reliable measure of time.
Yet it is difficult to believe that the rate of speciation
within the same genus, and involving in part the same
chromosomes, could have been roughly ten thousand
[ 287 ]
times as rapid in one period as in another. The difficulty
is rendered more acute by the necessity of assuming that
differentiation has been more rapid in the tetraploids
than in the diploids. Few serious students of evolution
will accept this premise.
In our opinion, the taxonomically distinct, wild, en-
demic, tetraploid cotton of Hawaii presents, for the
moment at least, an insuperable obstacle to the accep-
tance of the conclusions of Hutchinson, Silow and
Stephens. The case for the trans-Pacific, pre-Columbian
diffusion of Old World cultivated cottons is no better,
in our opinion, than the case for an Asiatic origin or pre-
Columbian diffusion of maize. To use the one as evidence
in support of the other, is to assume that two guesses
have, through some strange alchemy, a greater validity
than one.
Conclusion
We can find nothing in either the botanical or ethno-
graphic evidence presented by Stonor and Anderson on
Assamese maize to justify their conclusion that maize
must either have originated in Asia or been taken there
in pre-Columbian times. The maize itself is not unique,
since it resembles the living varieties of Colombia and
thus conforms to the general rule that all Old-World
maize has its counterparts somewhere in America. The
uses to which maize is put in Assam are exactly those
to which one would expect such a cereal to be put when
introduced into Asia, and there are no other special cir-
cumstances about its utilization, or the traditions con-
nected with it, which indicate a great antiquity in
Asia. The fact that maize, if introduced into Asia in
post-Columbian times, must have been rapidly accepted
by backward people, merely indicates that, like the po-
tato in Ireland, it met an acute and pressing need. Cer-
[ 288 ]
tainly there is nothing in the evidence which is in conflict
with the long-established and well-supported opinion
that maize is an American plant—one which has perhaps
been introduced into Asia twice: once in early post-
Columbian times from the west by a land route, and a
second time, perhaps somewhat later, when tobacco and
the potato were also introduced from the east by sea-
faring people. There is no factual evidence in conflict
with this simple and rational explanation; but there is
abundant evidence to support it.
The door is still wide open for hypotheses about pre-
Columbian culture diffusion between the Old World and
the New, and the problem is an extremely important one
which merits the most careful and critical attention on
the part of scholars in several fields. The problem is not
likely to be solved, however, by putting forward sweep-
ing and sensational conclusions which are based upon
inadequate and fragmentary evidence, especially when
these are all too likely to be seized upon by other imagi-
native writers who treat them as ‘‘evidence’’ or, worse
still, as ‘‘virtually unassailable proof’’ (Zelinsky, 1950).
Perhaps there has, indeed, been a pre-Columbian,
trans-Pacific diffusion of culture and perhaps maize has
been involved in it. To speculate upon this possibility
certainly can do no harm. But fancy ought not to be
confused with fact. The fact is, that, at the present time,
there is no tangible evidence of any kind — botanical,
archaeological, ethnographic, linguistic, ideographic,
pictorial or historical — of the existence of maize in any
part of the Old World before 1492. Until such evidence
is discovered, any case for pre-Columbian, trans- Pacific
diffusion must rest on evidence other than maize.
{ 289 ]
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