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
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.