CACAO AND ITS ALLIES 
A TAXONOMIC REVISION OF THE GENUS THEOBROMA 
Jose Cuatrecasas 
Introduction 
"Celebrem etiam per universam Americam multique usua 
fructum Cacao appellatum." 
Clusius, 1605. 
"Cacao nomen barbarum, quo rejecto Theobroma dicta est 
arbor, cum fructus basin sternat potioni delicatissimae, 
Baluberrimae, maxime nutrienti, chocolate mexicanis, Euro- 
paeis quondam folis Magnatis propriae (ffpotfta rwf 8&av, Vos 
Deos feci dixit Deus de imperantibus), licet num vilior 
fact a." 
Linnaeus, Hort. Cliff. 379. 1737. 
Theobroma, a genus of the family Sterculiaceae, is particularly 
noteworthy because one of its members is the popular "cacao tree" or 
"cocoa tree." The uses and cultivation of this outstanding tropical 
plant were developed in the western hemisphere by the Mayas in 
Central America a long time before Europeans arrived on the con- 
tinent. The now universally used name cacao is derived directly 
from the Nahuatl "cacahuatl" or "cacahoatl," just as the name of the 
popular drink, chocolate, is derived from "xocoafcl" or "chocoatl." 
The economic importance of cacao has given rise to great activity in 
several fields of development and research, especially in agronomy. 
Historians and anthropologists have also been very much interested 
in learning the role played by cacao in the economy and social 
relations of the early American populations. There exists today an 
extensive literature devoted to the many problems related to cacao. 
I saw cacao for the first time in Colombia in 1932, but became 
actually interested in the genus in 1939 and the years following, when 
I found cacao trees growing wild in the rain forests of the Amazonian 
basin. I was fascinated by the unique structure of the flowers of the 
cocoa tree, and its extraordinary fruit. My explorations from 1942 
379 
380 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
to 1947 at the service of the Government of the Valle del Cauca, 
mainly in the dense, humid forests of the Pacific coast of Colombia 
and western slopes of the Andes, offered me the opportunity to 
become better acquainted with wild species of Theobroma. I 
published the descriptions and illustrations of four new species 
found in that region, and at the same time gathered material for a 
monograph of the genus. Subsequently in Chicago and Washington 
I continued these studies, using the collections of the museums there 
and loans received from important European herbaria. The steadfast 
cooperation of V. M. Patino has been of great value to me in several 
respects, and his explorations have furnished two new species. 
Jorge Le6n from Turrialba sent me specimens of an outstanding new 
species. Collections sent to me for identification by the members of the 
English Colombian Cocoa Expedition (1952-1953) very much helped 
to broaden my knowledge of many of the species and their distribution. 
In 1954, I had the opportunity to study the important Theobroma 
collections at the British Museum (Natural History), London; the 
Royal Botanic Gardens, Kew; and the Museum d'Histoire Naturelle, 
Paris, where many Theobroma types are preserved. In 1961 (July 11- 
15) I was allowed to examine several European collections, at that 
time on loan at the Harvard Botanical Museum, which I felt desirable 
to see before publishing this revision. On my way to Colombia in 
1961 for a general collecting trip, I also visited the living Theobroma 
collection of the Imperial College of Agriculture in Trinidad where 
some 14 species and several hybrids and varieties are cultivated. I 
was thus able to supplement my data on the growing system and 
fruits of some of the species I had not seen living before. For the 
same purpose I visited the cacao stations of the Interam erican 
Institute of Agronomic Sciences at Turrialba, Costa Rica, 
This publication is restricted to the taxonomy of the genus. Many 
binomials (see the index) have been published in the past, and there 
is great confusion with regard to the names in the herbaria and 
literature. A critical revision was necessary in order to establish 
the validity of the species and to list synonymy. Although a complete 
study of the genus would require much more exploration, because 
of the great gaps existing in the herbarium collections, the present 
revision of all available materials seems justified. Three critical 
treatments of the genus have previously been published; Bernoulli 
(1869) recognized 18 species, and Schumann (1886) 11. Chevalier, in 
1946, acknowledged 13 species and a few subspecies; he united some 
species which should be held separate and on the other hand listed 
as different others which are actually synonyms. Twenty-two species 
are recognized in the present work. The number of species in the 
genus will probably increase in the future, because new explorations 
CUATRECASAS—CACAO AND ITS ALLIES 381 
in Central as well as South America will undoubtedly bring about 
the discovery of new ones. 
This revision is based on the classical method of comparative 
morphology. I have to a large extent used the structure of the fruit 
and the vegetative characters which I found were basic features in 
the definition and taxonomy of Theobroma, For a better under- 
standing of the genus, some new concepts are also contributed by 
G. Erdtman in pollen morphology (pp. 442-446), F. W. Cope in 
cytology and incompatibility in cacao (pp. 446-449), and by W. L, 
Stern in anatomy (pp. 439-442). These contributions may help in 
the understanding of the taxonomic problems of Theobroma, espe- 
cially those derived from cultivated varieties. In regard to T. cacao, 
the classification of the various cultivars is provisionally presented in 
a conservative way; an understanding of the innumerable existing 
forms of cacao will only be possible after long-term genetic research. 
Dr. Cope and Dr. Bartley in Trinidad are working in this direction. 
Dr. Soria in Turrialba, Costa Rica, is engaged at this time in an 
ambitious project of this kind, largely supported by the American 
Cocoa Research Institute of Washington. 
Due to the nature of this paper, the historical sketch is limited to 
the works which have contributed basic new data related to the 
taxonomy of Theobroma. The relationships with other groups and 
within the genus itself have so far as possible been treated objectively, 
with few hypothetical speculations. The specific descriptions are 
accompanied by original analytical drawings of the flowers, fruits, and 
leaves of almost all the species, and the illustrations, carefully 
supervised by the author, can be considered complementary to the 
written descriptions. Because of the relatively small number of 
existing collections, I think it useful to publish the information on 
herbarium specimens given by collectors; these data, except in 
special cases, have been translated into English whenever written 
in another language. However, the numerous herbarium collections 
of Theobroma cacao, mostly from cultivated plants, cannot be iden- 
tified as to the variety and are therefore not included in the text but 
simply listed in the index. For the citations of herbaria the abbre- 
viations of Lanjouw's "Index Herbariorurn" have been used. The 
abbreviation Photo F,M. is used to indicate the photographic series 
of the Chicago Natural History Museum. 
The artistic work for most of the illustrations has been carried out 
by the artists Christopher Reinecke, Maria Luisa Biganzoli, and 
Gil Cuatrecasas. Their work, consisting of about 35 plates, has 
been sponsored by the American Cocoa Research Institute; a few 
other plates had formerly been made by Gustavo Rojas, artist of the 
382 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Comisi6n Bot&nica del Valle, Colombia, and by Paula Gerard, 
Chicago. 
This revision has been done in the Department of Botany of the 
U.S. National Museum under a grant from the National Science 
Foundation. 
I wish to express my thanks to the directors and curators of the 
herbaria for their aid in the loan of collections and granting of work 
facilities, particularly to Dr. Albert C. Smith, Assistant Secretary of 
the Smithsonian Institution; to staff members of the Museum of 
Natural History, U.S. National Museum, who have checked the 
manuscript: Dr. Jason R. Swallen, head curator of botany, Dr. 
Lyman B. Smith, curator of phanerogams, and Conrad V. Morton, 
curator of ferns; to Dr. E. P. Imle, director of Research of the 
American Cocoa Research Institute; the Ministry of Agriculture of 
Colombia; to Mr. N, Y. Sandwith of the Royal Botanic Gardens, 
Kew, who helped on several occasions with important data and col- 
laborated in the typification of a few species; as well as to Mr. J. E. 
Dandy, of the British Museum, whose advice was of great help in 
the typification of T. cacao L. Likewise, I extend my thanks to 
Dr. W. Robyns of Brussels, Dr. H. Melchior of Berlin, Prof. H. 
Humbert, Prof. A. Aubreville, and Dr. A. Lourteig of the Museum 
National d'Histoire Naturelle, Paris, Drs. J. Pablo Leyva and 
A Fernandez Perez of the Institute de Ciencias Naturales, Bogota,; 
to my collaborators in Theobroma studies, Dr. Victor M. Manuel 
Patino, Ing. Humberto Guerrero, Dr. Ovidio Barros and Mr. Luis 
Willard, all of Colombia, and Dr. L. Aristeguieta, of the Institute 
Botanico, Venezuela. I further thank Drs. Cope and Bartley of the 
Imperial College of Trinidad both of whom were helpful in many 
ways, and Drs. Le6n and J. Soria of Turrialba, Costa Rica. 
The collections used for this revision are the following: 
Arnold Arboretum, Harvard University, Cambridge, Mass. (A). 
Botanical Museum, Harvard University, Cambridge, Mass. (AMES). 
Botanisches Museum, Willdenow Herbarium, Berlin (B). 
Bailey Hortorium, Ithaca (BH). 
British Museum (Natural History) London (BM). 
Jardin Botanique de l'Etat, Bruxclles (BR). 
Botanical Museum and Herbarium, Copenhagen (C). 
Institute de Ciencias Naturales, Bogota (COL). 
Edinburgh Royal Botanic Garden (E). 
Chicago Natural History Museum (F). 
Conservatoire et Jardin Botaniqucs, Genfeve (G). 
Goteborg Botaniska Tradgard, Goteborg (GB), 
Gray Herbarium, Harvard University, Cambridge, Mass. (GH). 
University of Glasgow, Dept. of Botany (GL). 
Systematisch-Geobotanischea Institut der Universitat Gdttingen 
(GOET). 
CUATRE CABAS—CACAO AND ITS ALLIES 383 
Staatsinstitut fur Allgemeine Botanik, Hamburg (HBG), 
Instituto Agron6mico do Norte, Belem do Pard (IAN). 
Royal Botanic Gardens, Kew, Surrey (K). 
Rijksherbarium, Leiden (L), 
Botanical Museum and Herbarium, Lund (LD). 
Komarov Botanical Institute of the Academy of Sciences, Leningrad 
(LE). 
Botanische Staatssammlung, Munich (M). 
Institute de Biologfa, Mexico (MEXU). 
Museu Goeldi de Historia Natural, Belem do Pard (MG). 
University of Michigan, University Herbarium, Ann Arbor (MICH), 
Missouri Botanical Garden, St. Louis (MO). 
New York Botanical Garden (NY). 
Museum d'Histoire Naturelle, Paris (P). 
Naturhistoriska Riksmuseum, Stockholm (S), 
Imperial College of Tropical Agriculture, Trinidad (TRIN). 
Botanical Museum and Herbarium, Utrecht (U). 
University of California, Berkeley (U C). 
U.S. National Herbarium (US). 
Facultad de Agronomfa del Valle, Cali, Colombia (VALLE). 
Institute Bot&nico, Venezuela (VEN). 
Naturhistorisches Museum, Wien (W). 
School of Forestry, Yale University, New Haven, Conn. (Y). 
Historical Sketch 
1605: First citation of cacao in botanical literature, by Charles de 
l'Ecluse (Clusius) in chapter XXVIII, of his Exoticorum libri decem, 
under the name Cacao fractus. It refers only to the fruit and gives 
a poor illustration of cacao seeds. "Celebrem etiam per universam 
Americam multique usus fructum Cacao appellatum." 
1623: K. Bauhin mentions for the first time in his books in his 
chapter "Amygdalus" the cocoa plant as "Amygdalis similis 
Guatimalensis Avellana Mexicana cujus fructum indigenae Cacao 
appellant," etc. (Pinax Th. Bot. 442). 
1630: The first prints of the Hernandez's Rerum Medicarum Novae 
Hispaniae Thesaurus appear in which are given descriptions of the 
cocoa tree under its Mexican name cacahoaquakuid and of four varieties 
called quaukcacahoatl, mecacahoatl, xockicacakoaU and llalcacahoatl 
which are distinguished by the fruits diminishing in size from the first 
to the last, presumably representing cultivars; the pods were called 
cacahoacentli and the useful seeds cacahoatl; he also mentions quauk- 
patachtli which undoubtedly refers to Theobroma bicolor. His illus- 
tration of the first (fig. 223) is clearly cacao Criollo. 
1658: W. Piso describes cocoa "De Arbore Cacavifera" and repeats 
the varieties cited by Hernandez in a long article on cocoa and 
chocolate. His drawing also represents the Criollo variety. 
384 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
1688: John Ray, in bia Historia Plantarum, gives much attention 
to cacao and its products in chapter VIII under the title: Cacao Ger. 
Cacao sive Cacavate Park. Cacao Americae sive Avellana Mezicana 
J. B. Amygdalae similis Guatimalensis C. B. The Cacao Tree. He 
explains that there are four kinds of cocoa trees and under the heading 
of Cacava guahuitl describes the tree, fruits, and seeds which he 
compares to almonds saying that they are white before ripening and 
red when fully ripe. 
1696: Plukenct, in his Almagcstum Botanicum, classifies cacao as 
an almond, under the name Arvor cacavifera Americana, the fruits of 
which (folliculi) contain some kind of almonds; in his plate 268, 
fig, 3, a leafy branch with a cacao fruit of the Criollo variety is 
represented. 
1696: Sloane lists Cacao in his catalog of Jamaican plants, with a 
long series of quotations from previous authors and travel writers. 
1700: The first taxonomic statement is made by Tournefort pub- 
lishing the genus Cacao: "Cacao est plantae genus auctore clariss. 
Plumerio" with a single species "speciem unicam novi." He gives a 
short description and drawings sent to him by Plumier. They dis- 
tinguish the 5 sepals, the strangulated petals, and a pistil surrounded 
by a laciniate girdle (staminodes) which develops into a ridged and 
pointed fruit filled with seeds. 
1705: Sibilla Merian gives an illustration of Surinam cacao which 
proves to represent clearly the Criollo type (26, t. 26). 
1710: Ray in his Methodus Plantarum copies Tournefort's descrip- 
tion and data to define Cacao; there are no changes in the 1733 edition. 
1725: Sloane publishes a long article on "The Cacao Tree" in his 
Voyage to Jamaica (vol. 2) giving the following botanical description 
(p. 15): "Out of the Body of the Tree, or Branch comes a very small 
Flower, standing on a half Inch long Footstalk, it is made up of 5 
Capsvlar Leaves, 5 crooked Petals, several Stamina, and a Stylus, of 
a very pale Purple color, after which follows the Fruit, which when 
ripe is as big as one's Fist, bigger in the Middle than at the Ends, 
which are pointed, it has some Salci and Asperities on its Outside, 
is for the most Part of a deep Purple colour, the Shell being about 
Half a Crown's thickness, and containing within it many Kernels 
of an oval Shape, each of which is as big as a Pistachio, Nut, having 
a thin Membrane without which is a mucilaginous Substance in which 
it lies. The Nuts themselves are made up of several parts like an 
Ox's Kidney, some Lines being visible on it before broken, and is 
hollow within, its Pulp is oyly and bitterish to the Taste, made up of 
many Striae, which tend from the Circumference to the Center." 
The plate 160 illustrates a leafy branch with oblong-ovoid fruits, 
10-ridged and strongly pointed; there are separate drawings of flowers 
CUATRECASAS—CACAO AND ITS ALLIES 385 
showing more or less clearly 5 sepals, some petals and (not well defined) 
4 or 5 staminode laciniae at the center, but no stamens; seeds, one 
isolated, some others together covered with pulp are also illustrated. 
The article gives much information about cultivation, varieties, 
geographical distribution, and trade in cocoa, and many authors are 
cited, 
17S7: Linnaeus (Genera Plantarum) introduces cacao into Classis 
18 of hia classification, in Polyadelphia pentandria giving it a new name 
Theobroma—meaning "food for the Gods"; the name Cacao given by 
Plumier and Tournefort was rejected by Linnaeus as "barbarous." 
By the International Code of Botanical Nomenclature, Linnaeus' 
name prevails. The new genus Theobroma was published almost 
at the same time in the Genera Plantarum and in Hortus Cliffortianus; 
Linnaeus described the flowers as having 5 stamens, 5 petals and 
staminodes (folioli nectarii), but only 3 sepals and with 5-celled 
anthers instead of 4. Linnaeus probably used dried specimens 
and annotations sent to him by Sloane for his description. He 
included in Theobroma two species: one with "foliis integerrimis," 
Cacao, the other with "foliis serratis," Guazuma, both also differen- 
tiated by their fruits. In the Genera Plantarum, he gives as the 
only synonyms and citations Tournefort for Cacao and Plumier for 
Quazuma, but in the Hortus Cliffortianus he quotes for Cacao: 
Clus.} Raj., Tourn., Sloane, MerHemPluk., and Bauh. But 
Linnaeus found out by himself, with the help of Sloane, the num- 
ber and kind of stamens typical of T. cacao. He writes in Hortus 
Cliffortianus: "Flores a nullo bene depicti, multo minus descripti 
sunt," and then: "Sloane mihi inspiciendi copiam fecit, videbatur 
structura exacte sequentis, ab aliis in universum omnibus diversis- 
sima." His original description of the stamens, given in the Genera 
Plantarum ("Filamenta subulata, longitudine nectarii, cui radiorum 
ins tar innata: singula apice quinquefida. Antheris in singulo stamine 
quinque, tectis petalo concavo"), was made on the basis of drawings 
or flowers sent to him by Sloane, for which reason the flowers of the 
Sloane herbarium have to be considered as the type of Linnaeus' 
description. But Linnaeus may have had very scanty material of 
the flowers, because he described the anthers as 5-celled instead of 
4-celled. 
1789: Weinmann writes extensively about cacao and chocolate and 
gives a plate (#77) which is inspired by Tournefort's illustration 
using very much imagination in painting it. Plate 278, devoted to 
Cacao minor, depicts a very deformed kind of pod. 
1739: Elizabeth Blackwell depicted cacao (pi. S7S) using Miller's 
specimens and suggestions. The fruit is figured as elongate, pointed 
386 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
and 10-ridged, although slightly verrucose; it represents the Criollo 
type. 
1741' Geoffroy in his De Vegetabilibus Kxoticis has a long article 
(XIX) De Cacao giving detailed descriptions of the tree, fruits, seeds, 
and their preparations. He calls the attention to the variability of 
the species in size or thickness of the different organs (especially the 
leaves and fruits). 
174^' Catesby publishes a magnificent colored plate of the cocoa 
tree in his Appendix to the Natural History of Carolina, etc. (p. 
6, pi. 6); the buds are colored red, the petals yellow, the staminodes 
red, and the fruits orange, obo void-oblong, 10-ridged and war ted, and 
very pointed; clearly it is the Criollo type. There is a long description 
of the tree with observations taken from D ampler; of the fruit it is 
said: "Fruit about the bigness of a swan's egg, but longer, more 
tapering, and ending in a point. The fruit hangs pendant, and when 
ripe, has a shell of a purple color, in substance somewhat like that of a 
pomegranate, and furrowed from end to end, containing in the middle 
many kernels of the size of acorns, inclosed in a mucilaginous sub- 
stance ..." In a later edition (1771) the color of the fruit was 
changed to dark violet. 
1749: Linnaeus, in his Materia Medica, includes Theobroma joliis 
integerrimis (p. 364) with the previous definition and classifica- 
tion; he attributes to it the qualities and virtues of pinguis, subamara, 
nutriens, aphrodisiaca, calefaciens. 
1758: Linnaeus gives a binomial name to cacao in his first edition 
of the Species Plantarum (p. 782): Theobroma Cacao, with the short 
specific diagnosis "joliis integerrimislie also names a second spe- 
cies Theobroma Guasuma with the diagnosis "joliis serratis." To the 
bibliographical citations are added: "Mat. Med. 364," "Geoffr. Mat. 
409," and llCatesb. car, 3. p. 6. t. 6." Inasmuch as the first edition 
of Linnaeus' Species Plantarum is the official beginning publication 
date for phanerogams by the Code of Nomenclature, Theobroma 
cacao receives here its official nomenclatural start, the generic name 
being in accordance with the diagnosis given in the corresponding 
edition (fifth) of Linnaeus' Genera Plantarum. 
1754: Linnaeus, in the fifth edition of Genera Plantarum, defines 
Theobroma and gives it the same classification (Polyadelphia pen - 
tandria) as in the first edition. Linnaeus did not improve his knowl- 
edge of the genus nor change his concepts of it in later works. In 
the third edition of Species Plantarum (1764), and also in the twelfth 
edition of System a Naturae (2: 508.1767) he keeps the same treatment 
of Theobroma as in his earlier publications. 
1754: The generic name Cacao is validated, according to the present 
Code of Nomenclature, by Miller in his fourth abridged edition of 
CUATRECASAS—CACAO AND ITS ALLIES 387 
"The Gardeners Dictionary." As in his popular sixth (1752) and eighth 
(1768) and other later editions of the Gardeners Dictionary, Miller 
publishes, without or with slight variations, a long article on "Cacao," 
"The Chocolate Nut." He explains that "This genus of plants was 
constituted by Father Plumier, who communicated the characters, 
which he had drawn in America, to Dr. Tournefort, who has inserted 
it in the Appendix of his Institutions. Dr. Linnaeus has joined this 
to the Guazuma of Plumier, under the title of Tkeobroma, but as the 
fruits of these plants are very different from each other, I shall keep 
them under different genera. We have but one species of this plant, 
which is Cacao." 
1763: Adanson separates Cacao (Tkeobroma) from Quazuma, but, 
while the latter is kept in the family "Les Tilleuls," Cacao is placed in 
the family "Les Pistachiers," side by side with Diosma, Triopteris, 
Acaju, Hugonia, etc., far away from its true relationships. 
1765: A disciple of Linnaeus, Antonius Hoffmann, presents to the 
Swedish Royal College of Medicine the first doctoral thesis ever pro- 
posed dealing with a Tkeobroma subject, "Potus Chocolatae." It is an 
excellent review of the knowledge about the composition and ways to 
prepare the chocolate at that time and the nutritional and medical 
importance of it. Hoffmann gives a more detailed description of the 
cacao plant and the fruit than did Linnaeus, probably inspired in 
Geoffroy; he writes: "Fructusmagnitudinem et figuram refert Melonis, 
sed verrucosus est, decem angulis instructus et superne acuminatus; 
dum maturescit, fit colore coccineus atque maculis variegatus flavis; 
intus con tine t nucleos circiter triginta, qui magnitudine divas aemul- 
antur ac pulpa obteguntur albida, subdulci et amaricante. Olei 
magna scatent hie nuclei copia, quod expressum vocatur Pinguedo de 
Cacao " (1769, p. 257.) 
1775: A disciple of Linnaeus, Jacobus Aim, in his doctoral disserta- 
tion Plantae Surinamenses republished ten years later in Amoenitates 
Academicae, emends the Linnaean description of the cacao flowers. 
He recognizes sterile stamens ("stamina alia 5, castrata") in what 
Linnaeus called "nectaria," alternate with the other "stamina fertilia 
solitaria"; he corrects Linnaeus also in seeing the calyx "pentaphyllus" 
and the anthers "quadriplici." 
1775: Aublet, in his explorations in French Guiana, found 
wild species of Tkeobroma, which he describes extensively in his 
Histoire des Plantes de la Guiane. He names them Cacao guianensis 
(pi. 275) and Cacao sylvestris (pi. 276); he quotes "cacao" as a Carib- 
bean name used. To the cultivated cacao, which he also found, he 
gives the new name Cacao sativa. It is unfortunate that Aublet mixed 
up elements of three species in describing his two new species; Cacao 
sylvestris was pictured from concordant parts of foliage and fruits 
388 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
which agree well with the species at present known as T. subincanum, 
but the flowers mentioned in his description were from T. cacao. 
Cacao guianensis was described and depicted from branches and 
foliage identical to Cacao sylvestris (= T. subincanum) and its flowers 
were taken from specimens of T. cacao; the fruit is the only different 
part, belonging to a third species. Aublet's descriptions and drawings 
are detailed, this being the first publication giving an accurate idea of 
the cacao flowers and of the fruits of some wild Theobroma. 
1785: Lamarck, in his famous Encyclop6die M6thodique, lists 
Cacao as belonging to the family of Cacaoyers characterized by 
hermaphrodite and complete flowers with 5 petals, 5 or 10 stamens 
and superior, usually 5-celled ovary; other genera of the family were: 
Ambroma, Guazuma, Ayenia, Buttneria, and Kleinhovia. He points 
out its close relation to Hermannia, Tilia, and the Malvaceae. He 
describes three species: Cacao sativa, C. sylvestris, and C> guianensis 
with the remarks mostly taken from Aublet, 
1789: A. L. Jussieu (Genera Plantarum) places Theobroma in Classis 
XIII, Ordo XIV (Malvaceae), of his system including it in his section 
V, (bis!) characterized by mixed fertile and sterile stamens connate 
at base; here Theobroma is associated with Pentapetes L., Abroma Jacq., 
Guazuma Plum., Melhania Forsk., Dombeya Cav., Assonia Cav., 
and Byttneria L. 
1791: Gaertner, in his remarkable book on fruits and seeds, gives a 
good description of cacao and drawings of its fruits and seeds, which 
represents a Criollo form; undoubtedly, Gaertner had at hand a dried 
specimen, for the section of the pod is drawn relatively thin and he 
describes it with "cortex sublignosus." The specific name used by 
Gaertner, Cacao minus, is a nomenclatural synonym of Theobroma 
cacao; Gaertner mentions Theobroma foliis integerrimis Linn., besides 
Sloane and Blackwell. 
1791: Schreber, in the eighth edition of Linnaeus' Genera 
Plan tarum, divides Classis XVIII in two subclassis, Decandria and 
Dodecandria. He places Theobroma in Decandria, a new concept, 
while Bubroma (a new name for Guazuma) and Abroma are brought 
into the Dodecandria. Schreber points out as differences between 
Theobroma and BvJbromay that the first has "laminae subrotundae 
acuminatae" and "antherae in singulo filamento duae," while Bubroma 
has "laminae semibifidae," "antherae in singulo fllamento tres," 
and "capsula non dehiscens muricata." 
1791: Gmelin, in the 13th edition of Linnaeus' Systema Naturae 
(vol. 2, p. 1151), includes Theobroma in Polyadelphia Decandria to- 
gether with the genus Abroma. He still lists T. Cacao and T. Guazuma 
as species of Theobroma, adding another: Theobroma guianensef a 
new combination based on Aublet's Cacao guianensis. 
CUATRECASAS—CACAO AND ITS ALLIES 389 
1796: Salisbury publishes Theobroma celtijolia, which is a synonym 
of Guazuma vlmijolia. 
1796; Lamarck (Tableau Encyclop&iique) publishes illustrations 
of Theobroma Cacao, Jig. 1 representing flowers, copied from Aublet's 
plate, Jig. 2 the smaller form of T. cacao illustrated by Gaertner, and 
Jig. 3 a, correct drawing of flowering and fruiting branches of a Criollo 
cacao (not T. guianense as stated). 
180B: Willdenow, in his edition of Linnaeus' Species Plantarum, 
follows the treatment of Schrcber, including Theobroma in Poly- 
adelphia Decandria and Bvhroma and Abroma in Dodecandria. Still 
only two species are considered: T. cacao and T. gujanensis. 
1806: Humboldt and Bonpland, in Plantae Aequinoctiales, a mag- 
nificent work, publish the first perfect botanical description of a species 
of Theobroma. It is supplemented with two plates illustrating leafy 
and flowering branches, fruits, seeds, and flowers of Theobroma bicolor, 
found by the authors cultivated in Colombia. The drawings show 
details of the embryo; the staminodes are wrongly figured as pointed 
instead of obtuse. 
1808: De Tussac, in Flora Antillarum, writes extensively on the 
cacao tree, its cultivation and uses, under the heading Cocas Theo- 
broma, giving "Le Cacaoyer Theobrome" as the French and "The 
Chocolate Tree" as the English name. There is a plate in folium 
(pi. XIII) showing illustrations of orange yellowish, 10-ridged fruits, 
attenuate at both ends, foliage, floral details, and the embryo. De 
Tussac is so enthusiastic about the use of chocolate that he says 
"le chocolat est au corps, ce que le caf6 est & l'6sprit" (p. 103). 
1811: Poiret, in the supplement to Lamarck's Encyclopedic, makes 
the new binomial Cacao bicolor, a combination based on the Humboldt 
and Bonpland species. 
1812: Stokes, in his Botanical Materia Medica, publishes a new 
name, Theobroma integerrima, for T. ca,cao L. 
1888: Kunth, in Humboldt, Bonpland, and Kunth, Nova 
Genera et Species Plantarum, gives a good description of the genus 
Theobroma and of the two species T. cacao and T. bicolor; the descrip- 
tion of the androecium is entirely correct: '' Filament a 10, basi in 
urceolum connata; quinque petalis opposita dianthera; quinque al- 
terna sterilia, lineari subulata. Antherae didymae biloculares, in 
petalorum cavitatae reconditae," Kunth gives a good description 
for the family Biittneriaceae R. Brown and for the five sections in 
which it is divided (Sterculiaceae, Biittneriaceae verae, Lasiopetaleae, 
Hermanniaceae and Dombeyaceae). Theobroma (jointly with Gua- 
zuma, Abroma, Glossostemon, Biittneria, Ayenia, and Commersonia) 
are placed in the Biittneriaceae verae which have the stamens 10-30. 
Filamenta magis minusve connata; quinque, laciniis calycinis opposita, 
390 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
anther is destituta, alius formae. Antherae didymae, longitudinaliter 
dehiscentes." 
18S4: De CandoIIe, in his Prodromus, accepts also family status 
for the Byttneriaceae (as Ordo XXVI) and follows the classification 
of Kunth, except for calling the sections divisions. In the tribe 
Byttneriaceae he includes the same seven genera, plus another doubt- 
ful one. In his treatment of Tkeobroma he includes five species, three 
already known, T. cacao, T. guianensis, and T. bicolor and two new 
ones, taken from the Mexican flora of Mocino and Sess6: T. angus- 
tifolia and T. ovatifolia. Mocino and Sess6 had written an unpub- 
lished Flora of Mexico supplemented by a series of illustrations, which 
were copied at Geneva by artists hired by De CandoIIe. 
1826: Sprengel, in the sixteenth edition of Linnaeus' System a Vege- 
tabilium, keeps Tkeobroma in Polyadelphia (Classis XVIII) and 
divides it in three genera: Tkeobroma with 2-antheriferous stamens 
and Bubroma and Abroma with 3-antheriferous stamens. He lists 
five species in Tkeobroma: T. cacao, T. bicolor Humb., T. speciosum 
W. herb., T. ovatifolium Sess., and T. guianense W. The genus 
Bubroma, considered synonymous with Guazuma Poir., comprises 
five species: B. Guazuma, B. tomentosum, B. polybotryon W., B. grandi- 
jlorum W. herb., and B. Invira W. Of these B. grandiflorum is 
actually a Tkeobroma. In Abroma, Sprengel included two species: 
A. augustum L. sup pi., and A. fastuosum Salisb. New Tkeobroma 
species in this publication are T. speciosum and B. grandiflorum. 
1827: Descourtilz, in his picturesque More MMicale des Antilles, 
dedicates much space to the description of the cacao tree "Cacaoyer 
cultivfi," to its origin, cultivation, varieties, uses, etc. Plate 266 
represents a leafy branch with flowers, seeds, and a fruit which is of 
the Criollo type (10-ridged, very warty, and acute). 
1828: Voigt publishes a description of a Tkeobroma guianensis 
without mention of Cacao guianerisis Aublet ("fol. acuminatis cor- 
datis sublobatis inaequaliter eroso-dentatis, subtus tomentosis, ram is 
petiolisque ferrugineo-hirtis, corymbo terminal!. Flores albi, parvi). 
According to some of the features given (corymbo terminali, flores 
albi), the plant described does not belong to Tkeobroma. 
1880: Sweet, in Hortus Brittanicus, listed four species under 
Tkeobroma mentioning a new binomial, T, caribaea, with no descrip- 
tion. It is undoubtedly a name for a form T. cacao. 
1830: Martius, who observed and collected many Theobromas 
on his trips throughout Brazil, says that more species may be found 
growing wild in tropical forests; he gives an account with short de- 
scriptions of the species found by him in Brazil, of which three are 
new species: Tkeobroma subincanum Mart., T, sylvestre Mart., and 
CUATRECASAS—CACAO AND ITS ALLIES 391 
T. microcarpum Mart.; the other species listed are T. cacao L. (= T. 
sativum Lam.), T. speciosum Willd. ?, and T, bicolor H. et B. 
1831: Don, in A General History of the Dichlamydeous Plants, 
places Theobroma in the family Byttncriaceae, tribe Byttnerieae 
DC. He gives good descriptions of these groups and of Theobroma, 
and short definitions for six species with some special attention to T. 
cacao. The other species listed are; T. guianensis, T. bicolor, T. 
angustifolia, T. ovatifolia, and T. sylvestris, the last being a new com- 
bination for Cacao sylvestris Aubl. 
1831: Martius, in his Reise in Brasilien (p. 1127), explains that 
the cocoa from Par A, and Rio Negro is of a lower, more bitter quality, 
because it comes more often from wild cacao trees than from culti- 
vated trees. He also says that he found T. bicolor growing wild in 
Barra do Rio Negro, in Manacurtl, and Yapura. 
1840: Endlicher, in his Genera Plant-arum, gives excellent descrip- 
tions for the Ordo CCXI Buttneriaccae and its six tribes, two of them 
being new: Eriolaenae and Philippodendrae; the other four are the 
same as those of Kunth and De Candolle, except for the Sterculiaceae 
which are treated as an order apart united with Bombacaceae and 
Helicteraceae. He includes in Bilttnerieae DC. the genera Rulingia, 
Commersonia, Abroma, Buttneria, Ayenia, Theobroma, and Guazuma. 
The genus Glossostemon is placed in the Dombeyaceae. 
1847: Dietrich, in his Synopsis Plan tar um, describes briefly nine 
species of Theobroma and gives a new name, Theobroma Martiana, for 
T. sylvestris Mart. 
1856: Karsten describes Theobroma glaucum, a new species from 
eastern Colombia. 
1861-1870: Baillon studies the development of the parts of the 
flower in T. cacao. The primordia of the sepals appear successively 
one after another, emerging above a common basal annulus. The 
petal primordia appear simultaneously and have the same aspect as in 
flowers of most other plants, but when they develop, there appears a 
strangulation which divides them into two articulated parts; the 
basilar parts are valvate, the upper parts contorted. The staminode 
primordia, opposite to the sepals, appear before those of the fertile 
stamens, which are opposite to the petals, and develop a simple fila- 
ment which divides into two, each branch having an anther whose 
two thecae become superimposed. The five primordia of the carpels 
opposite to petals have a half-moon shape; they become connate and 
develop alternate growths at the joints which are the primordia of the 
walls; these are centripetal and progressively divide the ovary into 
five cavities. Primordially, the placentation is parietal, becoming 
axile when the carpelar walls reach the axis; the ovules develop two 
integuments, become anatropous and placed in two rows in each 
680-69S—6d 2 
392 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
cavity, with the raphe, outside facing each other, horizontal. The 
embryo at first has ovate-orbicular entire, flat cotyledons; later these 
develop folds becoming corrugated. The initial transparent albumen 
is absorbed and at the end disappears. 
1862: Triana and Planchon, in Prodromus Florae Novogranatensia, 
list three species of Theobroma for Colombia (T. cacao, T. bicolor, and 
T. glauca) and three species of Herrania (H. pulckemma, H. albijlom, 
and H. laciniifolia). T. cacao is only given as cultivated, 
1862: Bentham and Hooker (Genera Plant arum 1: ix, 216), following 
basically the lines of De Candolle, although using other terms, classify 
Theobroma in the series Thalamiflorae, cohors VI Mai vales, ordo 
XXXII Sterculiaceae and tribus VI Buettnerieae. The chief charac- 
ters for the tribe are the hermaphrodite flowers, concave petal base, 
and anthers 1-3 alternating with staminodes; it is divided into two 
groups: *) the fertile stamens with 2-co anthers (Glossostemon, 
Abroma, Theobroma, Herraniat Guazuma), **) the fertile stamens with 
a single anther (Ayenia, Buettneriay llulingia, Commersonia). With 
the main characters of most of the genera already known, good defining 
descriptions are given for each of them. 
1869: Bernoulli attempts the first monograph of Theobroma. 
After a long time of field study in Central America and using the 
herbarium collections preserved at Berlin, Kew, and Munich, Ber- 
noulli became so well acquainted with the genus that he was able 
to draw a very good natural classification of it. The five sections 
established by him, based on flower and fruit characters, may be 
entirely kept today, strongly reinforced by other more recently known 
characters. When he started the study, there were only four or five 
species known besides T, cacao; and he writes, "Actually nobody knew 
these species, because the extremely short diagnoses of the old species 
were completely insufficient to determine them, for which reason there 
was the greatest confusion in the nomenclature in the herbaria that 
J had the opportunity to see." Bernoulli found that flowers and 
fruits give constant characters, whereas the leaves only give some 
secondary characters, especially the basal nervation; he also noticed 
that in some species there is great variation in the shape and pubes- 
cence of the leaves, with all transitions from one to another form 
when abundant material is compared. The basic characters used 
by Bernoulli for his system are: petal appendix or ligula sessile, 
subsessile or stipitate; staminodes erect or reflexed in bud, subulate, 
claviform, or petaloid; stamens 2-antheriferous or 3-antheriferous; 
fruit, when it was known; calyx 5- or 3-parted with narrow or broader 
lobes. With combinations of these features the following five sections 
(the descriptions here abridged) result. 
CUATRECASAS—CACAO AND ITS ALLIES 393 
1. Cacao: Petal ligule stipitate; staminodes erect, subulate; stamens 2- 
antheriferous; calyx 5-parted, the laciniae equal. Fruit ovate- 
oblong. 
2. Oreantkes: Fetal appendix subsessile; staminodes erect, subulate; sta- 
mens 3-antheriferous; calyx 5-parted, the laciniae equal. 
3. Rhytidocarpus: Petal appendix subsessile; staminodes erect, claviform; 
stamens 2-antheriferous; calyx 5-parted, the laciniae equal; fruit 
woody. 
4. TelmcUocarpus: Fetal ?; staminodes erect ?, linear-subulate with broad 
base; stamens 3-antheriferous; calyx 5-parted, the laciniae equal; 
fruit ovate, lacunose. 
5. Glossopetalum: Petal ligule stipitate; staminodes reflexed, petaloid; 
stamens 3-antheriferous; calyx irregularly 3-5-fid, "foliaceous"; 
fruit sublignose. 
Bernoulli describes 18 species, some of them extensively, others 
very briefly, according to the material he had at his disposal. Of 
these, 12 species are described as new. In sectio Cacao he includes 4 
species: T. cacao L. and 3 new ones: T. pentagona, "cacao lagarto" 
from Guatemala; T. leiocarpa, "cumacaeo" from cultivation in 
Guatemala; and T. saltzmanniana from Bahia, Brazil. In sectio 
Oreantkes he includes T. speciosa Willd. ex Spreng., and 2 new species: 
T, quinquenervia and T. spruceana, both from Brazil. Sectio Rhytido- 
carpus with one species, T. bicolor Humb. & Bonpl. (synonym, T. 
ovatijolia DC.), and, as doubtful, also T. glaum Karst. In sectio 
Telmatocarpus there is a single species, T. microcarpa Mart. In the 
sectio Glossopetalum, the largest, he describes in some detail T. 
angu&tijolia DC., T. subincana Mart., T. sylvestris Mart., and as new 
species T. macrantha from Brazil, T. jerruginea from Peru, T. obovata 
from Brazil, T. alba from British Guiana, and T. nitida from Brazil. 
About "Cacao guyanensis" Aublet, he writes "bleibt somit eine 
vollstandig ungewisse Art. Sie scheint auch von keinem weiteren 
Autor gesehen word en zu sein, sondern immer nur nach Aublet citiert 
zu Werden." Bernoulli quoted Herrania as a genus differing from 
Theobroma in the habit of the plant and in the 5-6-foliolate digitate 
leaves. Bernoulli's work is illustrated with several drawings. This 
monograph was published by the Swiss Society for Natural Sciences 
in its new serial of Memoires, vol. 24, no. 3, in 1871. However, a 
reprint was issued previously in pamphlet form in 1869, which is 
the effective date of publication. (See Fritzel, 1872; Sargent, 1912). 
C. G. Bernoulli was born in Basel, Switzerland, Jan. 24, 1834, and 
died in San Francisco, Calif., while returning home, May 18, 1878; 
he lived in Guatemala from 1858 until 1878 and collected plants and 
animals extensively in Central America. 
1873: Baillon publishes excellent comparative descriptions of the 
genus based on T. cacao in his Histoire des Plantes. He includes it 
in the family Malvaceae which includes the Sterculiaceae and Bomba- 
394 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
caceae; the family is divided into twelve series, the sixth being the 
Buettnerieae with 12 genera: Buettneria, Ayenm, Commersonia, 
Rulingia, Theobroma, Ilerrania, Guazuma, Scaphopetalum, Lepto- 
nychia, Abroma, Maxwcllia, and Glossostemon. The main characters 
given for the tribe are the usually hooded petal base, the fertile 
stamens opposite the petals, the staminodes alternate with the petals, 
the bilocular anthers (rarely 3-locular), the plurilocular ovary, and 
the capsular or carnose fruit. 
1882: D. Morris of Jamaica, in his well-known book on cultivation 
of cacao, after describing the cacao plant and fruit, publishes the 
first known classification of its varieties based upon the nomenclature 
of some of the best estates of Trinidad. He distinguishes two great 
classes (p. 12), the second being divided into several varieties: 
Class I. Cacao Criollo (Red). 
Class II. Cacao Forastero. 
var, a. Cundcamor verugoso amarillo (yellow), (rough yellow 
Cerasee) 
b. Cundeamor verugoso Colorado (red), (rough red Cerasee) 
c. Liso amarillo (yellow), (smooth yellow) 
d. Liso Colorado (red), (smooth red) 
e. Amelonado amarillo (yellow), (yellow Melon) 
/. Amelonado Colorado (red), (red Melon) 
g. Calabacillo amarillo (yellow), (yellow Calabash) 
k. Calabacillo Colorado (red), (red Calabash) 
This is the first valid publication in the nomenclature of the cultivars 
of Theobroma cacao. Morris goes on to say that before the "blast" 
or plague that almost exterminated cacao plantations in Jamaica 
before the end of the seventeenth century, the Criollo class was the 
only kind cultivated there. Since then, the Criollo has been entirely 
discarded for the hardier Forastero, the Criollo being "now chiefly 
confined to the mainland (Venezuela) where its yield, though small, 
is considered of great value" (p. 13). This assertion determines that 
the nomenclatural type of the Criollo cultivar is the Venezuelan 
Criollo. Of the Forastero varieties the best are the Cundeamor, the 
yellow kind being preferred for yielding a larger proportion of seeds. 
The seeds of Cundeamor "are mostly of the true almond shape— 
large, plump and full, of a pale crimson colour in the interior, and 
ferment easily." The Liso variety is closely allied to the former. 
The Amelonado is intermediate between Cundeamor and Calaba- 
cillo, and is considered as of good quality. The Calabacillo is the 
lowest quality and, Morris says, "never cultivated by a judicious 
planter; its fruits are small, the seeds flat, angular, intensely bitter 
and of dark crimson colour," 
1886: Schumann, in his Vergleichende Blii then morphologic, makes 
an accurate morphological analysis of eleven genera of Biittnerieae, 
CUATRECASAS—CACAO AND ITS ALLIES 395 
especially of Buttneria, Ayenia, Commersonia, Rvlingia, Guazuma, 
Theobroma, and Abroma, and tries to figure out the existing relation- 
ships between them and other Sterculiaceae genera. He concludes 
that the genera treated have a very close relationship and constitute 
a very natural group. Schumann sees in the Sterculiaceae two series, 
one of which is the main series (Hauptreihe), with a floral diagram 
C5, P5, Std5, A5) G5, where P.A.G. are opposite; these elements are 
either normally developed or modified by division or abortion; to 
this series belong as the leading tribe, the Buttnerieae, followed by 
Sterculieae, Helictereae, Lasiopetaleae and part of Hermannieae 
(Melochia, Dicarpidium, Waltheria). The other division of the Ster- 
culiaceae differs in having a floral diagram with the carpels opposite 
the sepals, this includes the Dombeyeae and Hermannia. He sees 
very close relationships between Buttnerieae and Lasiopetalae, which 
have to be artificially separated, and also between Biittneria and 
Ayenia, between Theobroma and Guazuma, and between the last and 
Scaphopeialum and Leptonyckia. There is no direct relationship 
between the two series of the Sterculiaceae, the Dombeyeae being 
close to the Malvaceae. Schumann does not see a way to explain this 
and decides not to draw any evolutionary hypotheses. 
Schumann describes the calyx of Theobroma as pentamerous or 
with three divisions, the sepals often concave and slightly cucullate, 
the petals sessile or shortly unguiculate. The inner glands of the 
receptacle are pluricellular and stipitate-globose or flagelliform; 
their function is unknown. The staminodes are more or less carnose. 
The ovules are in 2 rows in the ovary cells, but the seeds become 
uniseriate. The pulp of the seeds is mucilaginous and includes fine, 
curled, spiralish fibres. 
1886: Schumann, in the Flora Brasiliensis, gives a synopsis of 
Theobroma, describing with detail and accuracy its recorded species. 
But he does not follow the well-based system of Bernoulli. Schumann 
includes the genus Herrania in Theobroma as section Herrania, leaving 
the other species as section Eutheobroma. He uses the shape of the 
petal lamina and the staminodes and the number of anthers in a single 
stamen as secondary characters to classify the species using as the 
primary character the many-flowered or few-flowered inflorescence, a 
very vague feature in many instances. The keyed species in the 
first group are T. Cacao, T. bicolor, and T. speciosum, and in the 
second group, T. microcarpum, T. grandiflorum, T. subincanum, and 
T. angustijolium. Four species are listed outside of the key as 
"dubiae": T. glaucum Karst., T. silvestre Mart., T. Martii Schum., 
and T. album Bern. Two excellent illustrations are given for T. Cacao 
and T. grandifiorum. Schumann reduces to T. Cacao three Bernoulli 
species, T. leiocarpum, T. pentagonum, and T. salzmannianum, which 
396 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
he thinks were based on insignificant, variations of fruits or petals 
of this polymorphic species. Also he considers the flowers of Cacao 
guianensis Aublet as being T. Cacao, whereas the foliage and fruit is 
possibly T. subineanum. Bernoulli's T. quinquenervium and T. 
sprueeanum are reduced by Schumann to varieties of T. speeiosum. 
Schumann transfers Bubroma grandijlorum Willd. to Tkeobroma, citing 
as a synonym T. macranthum Bern. T. obovatum Kl. ex Bern, was 
not well known to Schumann, who makes it a synonym of T. subin- 
eanum. In short, the 18 species listed by Bernoulli are reduced by 
Schumann to 11, four of these being dubious to him. 
Schumann places Theobroma in the family Sterculiaceae, tribe IV 
Biittnerieae, characterized mainly by cucullate petals. This is divided 
into two sub tribes, corresponding exactly to the two groups made by 
Bentham and Hooker, but Schumann gives them names: a) Theo- 
brominae, defined as having 10 or 15 stamens and the base of petals 
cymbiform, and b) Buttnerinae, distinguished by having 5 stamens 
and the petal cucullus incurved at apex. Theobroma is the principal 
genus of the first, differing from Guazuma by its baccate fruit, lack of 
perisperm, and scarce, mucilaginous endosperm; Guazuma has lignose 
fruits, a developed perisperm, and lacks endosperm. 
1890: Schumann in his treatment of the Sterculiaceae for Engler 
and Prantl's Die Natiirlichen Pflanzenfamilien divides Theobroma into 
three sections: 1) Herrania, 2) Eutheobroma (2-antheriferous stamens) 
with T. cacao and T. bicolor, and 3) Bubroma (3-antheriferous stamens) 
with T. angustifolium, T. ovatifolium, T. grandijlorum, and T. subin- 
eanum. There are good illustrations. Schumann divides the family 
into eight tribes, Theobroma belonging to the fifth, Biittnerieae, 
characterized by hooded petals. It is divided into sub tribe Buttneri- 
nae with single stamens and Theobrominae with bundled stamens; 
the first includes Rulingiat Commersonia, Buttneria and Ayenia, and 
the second Glossostemon, Scaphopetalum, Leptonychia, Abroma, Theo- 
broma, and Guazuma. 
1890: G6mez de la Maza publishes Theobroma tomentosa, based on 
Guazuma tomentosa. 
1892: Hart modifies the Morris classification of the varieties of 
T. cacao, removing Calabacillo from the Forastero group and making 
with it a third class, as follows: 
Class I. Criollo or fine thin-skinned. 
1. var. a. Amarfllo 
2. var. b, Colorado 
Class II. Forastero or thick-skinned cacao. 
3. var. a. Cundeamor verugosa amarillo. 
4. var. 6. Cundeamor verugosa Colorado. 
5. var. c. Ordinary amarillo. 
6. var. d. Ordinary Colorado. 
CUATRECASAS—CACAO AND ITS ALLIES 397 
7. var. e. Amelonado amarillo. 
8. var. f. Amelonado Colorado. 
Class III. Calabacillo, or small-podded, thick, smooth-skinned, flat-beaned. 
9. var. a. Amarillo. 
10. var. 6. Colorado. 
Hart makes extensive and sound comments on the characteristics 
and qualities of each variety, saying, among other considerations, that 
" The finest cacao is by general consent admitted to be produced by 
the Criollo variety, and this is assumed to be identical or similar in 
character to that called the Caracas variety." (pag. 48). "The 
characteristics of the Criollo cacao are the thinness of its pod, its 
rounded beans and pale colour of the interior of the bean on section. 
The leaves of the tree are small when compared with the Forastero 
varieties, and the tree itself is not nearly so sturdy and thriving, and 
does not produce such regular and abundant crops as the Forastero 
and Calabacillo varieties. The skin of the bean itself is thinner, and 
the interior has but a small proportion of that bitter flavour which is 
characteristic of the unfermented bean of Forastero and especially 
that of Calabacillo. The flattest beans are those produced by pods 
of the Calabacillo type. The beans of Forastero are intermediate 
between these and the rounded form of the Criollo." (p. 51). Hart 
illustrates his important pioneer work with not too good illustrations 
of pods and its sections of Amelonado, Calabacillo, Forastero and 
Criollo, and with three diagrammatic sections of typical seeds of Criollo, 
Forastero and Calabacillo. He adds, however, that there will be 
found intermediate forms hardly reconcilable with any of the figures, 
so that "these are to be taken as representative only of the typical 
varieties with some latitude." 
Hart's illustrations are not representative of the concepts for which 
the same nomenclature is generally used outside Trinidad: For 
instance, the figure given as Criollo represents the type Cundeamor, 
and the models he used to illustrate Calabacillo were not well selected. 
These facts explain the comments made some years later by Preuss 
about Hart's somewhat confusing nomenclatural concepts. 
1898: John Donnell Smith describes a new Theobroma with yellow 
flowers and smooth cylindrical pods which Pittier and Tonduz dis- 
covered in the mountains of Costa Rica, T. simiarum. 
1899: Jumelle publishes an excellent monographic compilation on 
cacao with a long chapter devoted to the botany following an inter- 
esting account of its history. Disregarding the work of Bernoulli, 
Jumelle follows Schumann in his treatment of the Sterculiaceae in 
the Flora Brasiliensis, declaring that "C'est certainement l'6tude la 
plus complete et la plus consciencieuse qui ait 6t6 faite sur Ies Theo- 
broma.'1 Then he gives in French Schumann's complete key for seven 
398 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
species. He recognizes that several species Jhave been published 
before, based on weak characters, probably due to variations caused 
by cultivation or changes of soil or climate. Jumelle devotes much 
attention to the description of T. cacao and its different varieties. 
He follows in this the works, experiences, and classification of Hart. 
Chapters with descriptions and accounts are also devoted to T. bieolor 
(and separately T. ovatijolium), T. angustifolium, T. subincanum) 
T. grandiftorum, T, speciosum, T. microcarpum, and T. glaucum. 
Short notes are given to T. sylvestre Mart., T. Martii Schum., and 
T. album Bernoulli. Although excellent as a compilation, Jumelle's 
publication offers nothing original with regard to the systematics of 
Tkeobroma; its many illustrations were taken from Bernoulli, 
Schumann, and Hart. 
1899: De Wildeman publishes a new binomial, Tkeobroma Kalagua, 
with a description and illustrations of leaves, flowers, and fruit. 
Unfortunately, this "new species" was based on separate elements 
belonging to different trees and different species from unknown 
localities, sent to de Wildeman by a Mr. Ch. Patin, at that time 
vice consul of Belgium in Panama. Although the specimens used 
for the description were claimed by Patin as having been collected 
from a single tree, Patin himself recognized later that they came from 
different trees and that the fruits were from T. simiarum. At that 
time, Panama was part of Colombia and that is the reason why the 
new species and the specimens were labeled as from Colombia. 
1900: In a second work, Hart makes no alterations in his 1892 
classification, stating that "after the lapse of some years I still see 
no necessity to revise the list." But he extended his comments on 
characteristics, variations, and properties of the cacao varieties with 
new considerations, as for instance: "Calabacillo is certainly as far 
removed from Forastero, as Forastero is from Criollo, as seen in 
plantations of the present day, when every intermediate form from 
Criollo down to Calabacillo can be seen linking the whole in one 
continuous chain of varieties. To properly classify Cacao, we must 
first know what the originals were like, and it is clear that at the 
present time, it is hard to decide exactly what were the forms assumed 
by the older types of Cacao fruit. There is an apparent consensus 
of opinion however which points to the thin-skinned and bottle- 
necked variety as the original Criollo (Spanish for Creole), and this 
is quite confirmed by the Criollo being discovered in the virgin forest 
of an uncultivated part of Trinidad" (p. 52). 
The Criollo as well as the cacao of Java and Ceylon, the Criollo 
of Central America and T. pentagonum have the seeds white or almost 
white inside. The best quality Forasteros have the seeds slightly 
violet, and the Calabacillo strongly colored. He adds that the best 
CTJATRECASAS—CACAO AND ITS ALLIES 399 
qualities of fruits of Venezuela (e.g., from Ocumare) are distinguished 
by the lightness of the seeds, and their shape, although the pod might 
belong to the type of Forastero. In Trinidad, in the Estates where 
certain strains of cacao from the continent had been introduced, we 
find the finest qualities of Forastero. The illustrations, with a 
similar nomenclature, are less clear than those of first edition, 
1901: Paul Preuss publishes the first important field report ever 
written with keen observations on the varieties of cacao, conditions 
of cultivation, conditions and ways of preparation, and qualities of 
the products in several countries of South and Central America. It 
is the report of his trip made during 1899 and 1900, in order to obtain 
information on tropical crops for the Colonial-Economic Committee 
of the German government. From Surinam he names three varieties 
of T. cacao as cultivated: "Surinam/' also called "Porcelaine" (cor- 
responding to the Amelonado of Trinidad), "Aligator" (the Cunde- 
amor of Venezuela and Trinidad), and "Caracas" (similar to Carupano 
or Forastero from Trinidad), He also observed mixed intermediate 
forms. From Trinidad and Grenada, where he saw extensive planta- 
tions, he describes several cultivars: "Amelonado," "Calabacillo," 
"Sangre de toro," "Forastero," "Criollo"; he mentions that Hart 
calls Criollo what is called Forastero in Venezuela and that Hart's 
Forastero is the Venezuelan Criollo. But according to Preuss, the 
formerly general use of the name Forastero in Trinidad applied to 
cacaos other than Criollo. At present, Forastero has the same 
meaning as Trinitario, which might have been introduced from the 
Venezuelan eastern coast or from the Orinoco region, after the earlier 
Trinidad plantations (all of the Criollo type) had been destroyed by 
some kind of disease. Preuss goes on to describe the three main 
Trinidad varieties using the concepts of Hart: "Forastero," "Amelo- 
nado," and "Calabacillo." He compares the yellowish variety of 
Amelonado to the Guayaquil cacao and the elongated form of Foras- 
tero to the Colombian cacao. Of Venezuela, Preuss says that it 
is the classic land of Criollo, producing large seeds with thin shell 
of the best quality. He distinguishes two main cacao varieties in 
Venezuela: 1) Cacao Criollo, 2) Cacao Trinitario. Preuss clearly 
distinguishes seven types, but there are also many intermediate 
forms. 
1. Angoleta. 
2. Cundeamor (Cundeamor legitimo, with red shell and Cundeamor amarillo, 
with yellow shell). 
3. Carupano legitimo (Carupano grande, and Carupano mestizo, red with 
yellow). 
4. Carupano parcho (yellow fruit). 
5. Carupano Taparito (yellow or brownish). 
6. Sambito (red or yellow, short, thick, rather smooth fruit). 
400 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
7. Trinitario amargo or "Coj6n de toro" (red or red brownish, smooth, blunt 
or shortly attenuate pointed fruit). 
He describes the Criollo tree and mentions its three varieties: "Criollo 
legitimo" (the best quality) with deep red, "Criollo amarillo" with 
yellow, and "Criollo mestizo" with yellow and red shells. The 
seeds are white violet in the first and white in the second. Preuss sees 
in Ecuador a great uniformity in the cacao fruits as he had observed 
in no other country. They are of the Amelonado type, the same type 
being cultivated in the Cameroons, St. ThornG, Grenada, and Surinam; 
the Ecuadorian type has thicker shells. He mentions four varieties: 
"Arriba," "Balao," "Machala," and "Bahia." He says that cacao 
is frequent in the underlayer of the rain forests in Ecuador and that 
these wild trees, which cannot be distinguished from those of the 
plantations, also produce good fruits, which have very thick shells 
and roundish seeds in section. Preuss sees in this wild cacao the 
origin of the widespread plantations of the variety Amelonado (p. 
247). Theobroma bicolor is also mentioned as a wild cacao ("cacao 
bianco," "bacao") in Ecuador. 
From Central America he describes the native variety "Cacao 
Lagarto" (Alligator), the other native variety Criollo and two others 
introduced, the "Cauca" from Colombia, and "Trinitario" from 
Trinidad; in this Trinitario he distinguishes the three types Forastero, 
Amelonado and Calabacillo. There are intermediate forms between 
Lagarto and Criollo "Cacao del pais" difficult to separate one from 
another; they have red and yellow pods with a thin shell and white- 
violet or white cotyledons. The seeds of Nicaraguan Criollo are the 
largest of all varieties and of the best quality known. Mention is 
made and illustrations are given of T. bicolor, T. angustijolium, and 
T. pentagonum from Central America. Also drawings of fruits and 
seeds of five varieties of T. cacao (Nicaraguan and Venezuelan Criollo, 
Calabacillo, "Cundeamor Legitimo," and "Curupano grande") are 
reproduced. 
190$: De Wildeman gives a taxonomic compilation of Theobroma 
in his book on tropical cultivated plants. Following Schumann he 
divides the genus in Ilcrrania, Eutkeobroma, and Bubroma and uses 
Schumann's key for the species (as in the Flora Brasiliensis), but adds 
another species, T. simiarum Donn. Smith, and accepts T. 'pentagonum, 
as different from T. cacao; altogether, nine keyed species and four 
doubtful additional ones, as by Schumann, besides Herranias, are 
treated. For each species, summarized descriptions, native names 
and uses, geographic location, and miscellaneous comments, are 
given. Speaking of T. simiarum he writes: "Tl faut rapporter en 
partie & cette esp6ce la plante que nous avons dccrite, en 1899, sous le 
nom de T. kalagua, dont la description avait 6t6 fait sur de feuilles, 
CUATRECASAS—CACAO AND ITS ALLIES 401 
des lleurs et des fruits ne provenant pas de la m&me plante." Some 
attention is given to T cacao and the differences between its seeds and 
those of Herrania mariae, based on anatomical sections and gross 
chemical analysis made by Heim. The varieties of Tkeobroma cacao 
are reduced to ten "series" distributed in three "grand" groups— 
I) Criollo with two varieties: 1) amarillo, 2) Colorado; II) Forastero 
with varieties: 3) cundeamor verrugosa amarillo, 4) cundeamor 
verrugosa Colorado, 5) amarillo, 6) Colorado, 7) amelonado amarillo, 
8) amelonado Colorado; III) Calabacillo with varieties: 9) amarillo, 
and 10) Colorado. 
1904: Lignier, in a list of the Caen City Herbarium, mentions 
Theobroma sativa, as a Sagot collection from French Guiana without 
further comment on the author or synonym; it probably refers to 
Cacao sativa Aubl. 
1904: Huber, a botanist at the Goeldi Museum, Bel£m do Par A, 
inaugurates a new epoch by publishing direct observations on the 
botany, ecology, and location of species of Tkeobroma. Huber found 
spontaneous and subspontaneous trees of T. caca,o in several places of 
Amazonia and believes that it may really be indigenous in the forests 
of the Alto Purtis Rio, Rio Ucayali, and other places down to San- 
tar&n and Obidos. 
1906: Huber extends his explanations on the indigenism of T, 
cacao on the alluvian soils of the Rio Alto Pur us, in the inundatable 
forests around the mouth of Rio Acre, and along the rivers Ucayali, 
Japur6, Juru6, and Madeira. Among observations on other species 
(T. microcarpum, T. speciosum, T, obovatum (as T. sylvestre), T. 
subincanum, and T. bicolor) he shortly describes a new species from 
Peru, Theobroma sinuosum Pa von. 
1906: Huber publishes a variety coriaceum of T. speciosum found 
in Brazil. At the same time, he mentions having found T. cacao 
growing wild in a forest near the Canchahuaya Lake. 
1908: Chevalier, in his extensive studies on cacao crops in western 
Africa, besides detailed descriptions of the cocoa tree and its growth 
and ecology in Africa, mentions its variations and describes a new 
species, Tkeobroma sphaerocarpa, distinguished by its globose, almost 
smooth fruit 9 to 11 cm. in diameter, which has been cultivated for 
a long time on the island of Sao Tom6. 
1911: Hart, in the introduction of his book on cacao, gives an 
excellent account of the varieties of the cultivated cacao, discussing 
the external features of the chief different types and the qualities of 
their crops: 
"The species known as Theobroma cacao, covers innumerable varie- 
ties or forms, differing in shape of pods, in size and vitality of trees, 
in bearing capacity, and in colour, shape and quality of the bean. 
402 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
The many names under which varieties of this tree {Theobroma 
cacao) are known, do not constitute species, but must be merely con- 
sidered as varieties of one species. These varieties probably owe 
their origin to seed variation and cross-breeding, together with the 
local influence of soil and climate, but it would serve no useful purpose 
to record the names by which they are known, as these differ in each 
district, in each plantation, and in each country where they are 
grown" (p. 2). 
"We have therefore a classification under Theobroma cacao carrying 
fourteen types under three classes, but it must be understood that 
these are separated by no definite margin, and that intermediate 
forms will be found on estates showing every conceivable form of 
variation" (p. 3). 
Hart's classification here amplifies his initial one of 1892, mainly 
by broadening the concept of Criollo, in which class he includes types 
known under this name outside Trinidad a long time before. 
Theobroma cacao 
Class I. Criollo 
Trinidad Criollo. 
(1) Var. a. Amarillo=Yellow, thin-skinned, bottle-necked. 
(2) Var. b. Colorado = Red, thin-skinned, bottle-necked. 
Venezuelan Criollo. 
(3) Var. o. Amarillo=Yellow, thick-skinned, high-shouldered, some- 
times pointed. 
(4) Var. b. Colorado=Red, thick-skinned, high-shouldered, some- 
times pointed. 
Nicaraguan Criollo. 
Thick-skinned, high-shouldered, and 
(5) Var. a. Amarillo—Yellow 
(6) Var. b. Colorado—Red 
very large beans with light-coloured 
interior. 
Class II. Forastero 
(7) Var. a. Cundeamor verugoso Amarillo = Yellow-warted. 
(8) Var. b. Cundeamor verugoso Colorado = Red-war ted. 
(9) Var. c. Ordinary, or typical Amarillo = Yellow Forastero. 
(10) Var. d. Ordinary, or typical Colorado = Red Forastero. 
(11) Var. e. Amelonado Amarillo = Yellow, melon shaped. 
(12) Var. /. Amelonado Colorado = Red, melon shaped. 
Class III. Calabacillo 
(13) Var. a. Amarillo = Yellow 1 Calabacillo, flat-beaned, smooth, thin- 
(14) Var. b. Colorado = Red / or thick-skinned, and small pods. 
Theobroma pentagona 
(15) Theobroma pentagona=Alligator cacao. Has yellow, mueh- 
warted pods, with five distinctly 
raised ribs, and large beans, having 
white or light-coloured interior. 
CUATRECASAS—CACAO AND ITS ALLIES 403 
Hart characterizes the Criollo varieties by their light-colored seeds, 
the high quality of the cured product, and the less vigorous growth. 
The Forastero has light to dark purple seeds in large rough-ridged 
pods; it is rather variable and is a strong grower. Calabacillo is an 
inferior but stronger growing tree, having ovoid pods with thin, 
solid, dark-colored seeds. (tT. pentagona has the largest seeds of 
any known species." "In general, however, it is hard to say where 
one form begins and another ends." In fact in most countries, 
cacao "consists of a heterogeneous mixture of cross-bred varieties of 
one species (T. Cacao) though of late years it is thought possible that 
the common species may have become hybridized with T. pentagona " 
1914 and 1932: Van Hall publishes his well-known book on cacao 
which since then has been a textbook and main source of information 
for cacao growers and agronomists. In his botanical chapter he 
presents some nonoriginal information on the noncultivated Theo- 
bromas. He gives excellent comparative descriptions and evaluations 
of the most important variations and forms of cultivated cacao. 
He follows Morris in recognizing two groups: 1. Criollo; 2. Foras- 
tero. He describes seven types or sub varieties of the first: the 
Venezuelan, the Ceylonese, the Javan, the Samoan, the Madagas- 
car! an, the Nicaraguan, and the Surinam Criollos. The subvarieties 
of Forastero described are Angoleta, Cundeamor, Amelonado, and 
Calabacillo. Theobroma spkaerocarpa Che v. is considered a mere 
form of Calabacillo. In 1932, van Hall introduces a new name 
T. aspera, transferred from Herrania. 
1914* Pit tier publishes two new species of Theobroma from Panama: 
T. bernouillii 1 discovered by him in the forests of the Colon province, 
and T. purpureum which belongs to Herrania. 
1915: Cook, after a thorough morphological field study of T. bicolor 
and T. cacao, especially of the branching system and the flower and 
inflorescence structure, decides that they belong to different genera, 
and publishes a new monotypic genus Tribroma with one species, T. 
bicolor; it mainly differs by its clusters of three branches and the woody 
pericarp in contrast to the 5-branching clusters and fleshy pericarp of 
T. cacao. Although all observations by Cook are very sound, he does 
not compare T. bicolor with other species of Theobroma besides cacao. 
1916: Cook publishes in detail and with illustrations the results of 
his studies on growth and dimorphism of branches and leaves of T. 
bicolor and T. cacao. He explains the sympodial structure of the 
Theobroma stem, the verticillate primary branching, the formation 
and succession of upright lateral shoots, which continuing the main 
stem, bring the clusters into a lateral position and the alternate 
i Originally bo misspelled, although the correct name would have been ubtrnwUU,** 
404 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
branching of lateral branches. Cook describes the two types of leaves: 
the long petiolate symmetrical leaves of the seedlings and upright 
shoots and the bilateral, asymmetrical, dorsiventral, short-petiolate 
leaves of the lateral branches. The structure of inflorescences and 
flowers also are studied. 
1918: Stahel publishes a precise, illustrated, morphological study 
of the structure of the inflorescences of T. cacao and T. bicolor. These 
inflorescences are always formed in the axil of the subtending bract of 
a lateral, non- or short-developing bud. The inflorescence in T. cacao 
consists of a cincinnous main axis with many short internodes and a 
few dichasial final branchlets. In T. bicolor, the main cincinnous axis 
has a few, long internodes, and the lateral branching is dichasial. The 
peduncles have a bilateral, the pedicels a radial structure. 
1921: Benoist publishes a new species of Theobroma from French 
Guiana found by himself, T, velutinum. It is characterized by large 
leaves, velutinous beneath, and by ellipsoid, 5-ridged, velutinous pods. 
1023: Standley recognizes three spontaneous species for the Mexican 
flora: T. cacao, T. augustifolium, and T. bicolor, and gives interesting 
historical, geographical, and economic data. 
1925: Ducke publishes firsthand new botanical and ecological data 
on several Brazilian species. 
1925: Pittier presents his interesting new theory that all existing 
forms of cultivated cacao are the result of hybridization between two 
initial species: 1) Theobroma cacao L., with elongate, claviform, rugose, 
10-ridged pods, containing large, ovoid, white or slightly yellowish 
seeds, and 2) T. leiocarpum Bernoulli, with more or less rounded, 
smooth, slightly 5-ridged pods, with flattened, more or less triangular 
and dark purplish seeds. The first type is the one commonly known 
as "Cacao dulce" or "Cacao criollo," the second is commonly known 
as Calabacillo or Trinitario and in Guatemala as "Cumacaco." T. 
sphaerocarpum Che v. from Sao Tome is a hybrid, retrogressive to T. 
leiocarpum. The two typical forms are united through an unlimited 
number of intermediate hybrid forms the characteristics of which are 
very variable; he says that the present nomenclature of varieties 
should be abandoned. 
1925: Chevalier, in his "Observations" to the preceding work of 
Pittier, accepts basically Pittier's theory, and says that T. sphaero- 
carpa would be the extreme form of the series with smooth fruits and 
that T. leiocarpa is a hybrid between T. cacao and T. sphaerocarpa. 
1926: Pittier answers Chevalier saying that he has never found T. 
sphaerocarpum in Central America, but that he has found growing 
wild in Costa Rica and Panama forms almost identical to T. leiocarpum 
Bernoulli. He recognizes that the forms with small, round pods often 
found in the region of Barlovento (Venezuela) are very close to T. 
CUATRECASAS—CACAO AND ITS ALLIES 405 
sphaerocarpum. Both original species are still found in pure, typical 
form and it can be said that in most plantations, next to the "primitive 
species" a number of hybrids are found. "We have still in Venezuela 
cacao plantations with absolute domination of the Criollo type, as, for 
instance, it happens at Caruao and Chuao." There can be observed 
over great extensions of land, tree after tree with elongated, claviform, 
pointed fruits, which may be reddish or yellow, with rounded seeds and 
almost white and insipid cotyledons, Pittier adds that at the time 
Linn6 described T. cacao, the Criollo was the dominant form in 
cultivation. 
1930: Myers reports about his exploration of the upper Mamaboen 
Creek, a tributary of the Coppename River, where wild cacao 
was discovered by Stahel ten years earlier. This place is located in 
the middle of the Surinam rain forests, 40-50 kms. away from the last 
small Indian village. Abundant trees of cacao in a wild state were 
found in the lower tree-layer of the forest, under conditions of dense 
shade and high humidity; most of the cacao trees were found on the 
flooded margins of the river; few were found on higher ground. The 
trees were 10 to 25 feet high, and abundantly fruiting; the ripe pods 
were bright, light yellow in color, and almost smooth (with little indi- 
cation of longitudinal ribs) and with 40 to 50 seeds with deep violet 
cotyledons. Myers adds that the fruits were of the Amelonado- 
Forastero type. He also comments on information about wild trees in 
other parts of Surinam, British Guiana, Brazil, and elsewhere. 
1930: Pittier publishes an abridged key to classify the known species 
of Theobroma, including Herrania. Following Schumann in Die 
Natiirlichen Pflanzenfamilien he divides it into three sections, the first 
being Herrania. The other two sections, defined by diantheriferous 
(Sect. Eutheobroma) and triantheriferous stamens (Sect. Bubroma), 
comprise 13 species, which are distributed in subsections corresponding 
to the Bernoulli sections. In Eutheobroma he includes subsect. Cacao 
with T. cacao, T. leiocarpum, and T. pentagonum and subsect. Rkyti- 
docarpus with T. bicolor and T. BemouiUii. In Bubroma there are 
subsect. Telmatocarpus with only T. microcarpum, subsect. Oreanthes 
with T. spruceanum, T. specios-um, and T. simiarum and subsect. 
Olossopetalum with T. angustifolium, T. grandijlorum, T. subincanum, 
and T. sylvestre. The characters given in the key are few and not 
always the most typical or correct, some of the species being wrongly 
placed (T. Bernov/Ulii, T. simiarum). The binomials T. glaucumt 
T. martii and T. album are considered dubious. Pittier refers to 
having received from Cook photographs of two forms of cacao culti- 
vated in Peru with small fruits which might be different species; one 
has almost spherical, rugose pods called locally "cacao chuncho." 
Pittier says that, with the exception of T. leiocarpum and T. cacao 
406 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
(respectively Calabacillo and Criollo), there do not exist stable forms 
of cultivated cacao. Forastero from Trinidad stands between these 
original species and can be considered as the result of their crossing. 
The Trinidad Forastero seems to be quite different from Venezuelan 
and Central American Forastero and probably from the Old World 
Forastero. He also says that it does not make sense to talk about 
the origin of Forastero, because its forms appear wherever the parental 
species are present and successive crossings produce constant varia- 
tions. Commenting on the finding of Myers in Surinam, Pittier says 
that the wild cacao of the Mamabocn valley belongs to T. leiocarpum. 
The same opinion is expressed in connection with Schomburgk's 
spontaneous cacao found in the Rio Branco valley and numerous 
references about wild cacao throughout the Venezuelan Guayana. 
The cacao with tapering, pointed, ridged and rugose pods with white 
cotyledons {T, cacao) has really never been found indigenous to the 
east of the Panama isthmus. Pittier found it at the isthmus but says 
that the origin of this species is to be sought towards the north, in the 
Soconuzco, Chiapas, and Tabasco regions, where the Criollo type 
finds the best conditions for its development. Conversely, the planta- 
tions of Calabacillo do not go much further west than Costa Rica 
where it was recently introduced. Here are to be found the western- 
most stations of T. leiocarpum, although the type came from planta- 
tions in Guatemala. 
1981: Mildbraed publishes a new species T, tessmannii, found by 
Tessmann in eastern Peru. He relates it to T. ferrugineum Bernoulli, 
from which it differs especially by the long, soft tomentum of the 
leaves beneath. 
1932: Cheesman, who had been conducting research on cacao corps 
for many years, presents a significant account of the economic botany 
of Theobroma. He believes that it is unlikely that any of the now 
uncultivated species has any direct economic significance. He speaks 
about the cultivated species, those of section Cacao, as interfertile 
species {T. cacao, T. leiocarpum, T, pentagonum). Cheesman says 
that "the taxonomic status of the group of forms at present included 
under the collective term T, cacao can only be determined by pro- 
longed research, including genetic, and possibly also cytological 
studies." Cheesman agrees with Pittier in considering that more 
than one species contributed to build the "cacao complex," adding 
that this idea provides the most helpful way of regarding the ex- 
traordinary variation exhibited by the crop. His discussion on the 
history, characters, and merits of the varieties is illuminating in many 
respects, especially when trying to understand their possible origin. 
In practice, he follows, with slight modification, van Hall's classifica- 
CUATRECASAS—CACAO AND ITS ALLIES 407 
tion which he says "is a compromise between a natural and an artificial 
system." 
1932: Pit tier insists in his viewpoints on the origin of the cultivated 
cacao. Finding that T. cacao L. is not well typified by a known 
variety, Pittier decides to abandon this name and to substitute for it 
T. sapidum, but no specific description is given and a type specimen 
is not indicated. The Calabacillo seems to have a stronger fertilizing 
power than the Criollo which explains why few trees of the first suffice 
to alter the plantations of the second. 
1933: Ciferri makes a very detailed, critical study of the cultivated 
cacaos of Santo Domingo giving a thorough classification with defini- 
tion, descriptions, and illustrations of their numerous types. Mor- 
phological, taxonomic, historical, and economic comments are given. 
Ciferri follows the principles of Pittier in accepting the theory that the 
majority of the cultivated cacaos are hybrids of two initial types, 
but he considers these two types varieties instead of species. Ciferri 
gives the system formal nomenclatural status by publishing the fol- 
lowing new varieties: T, cacao L. emend. var. typica Ciferri; T. cacao L. 
emend, var. leiocarpa (Bernoulli) Ciferri, and T. cacao var. typica X 
T. cacao var. leiocarpa = Forasteros. "I cacao denominati global- 
men te "Forasteros" sarabbero dunque, secondo l'idea di Pittier, che 
moi adottiano in pleno, i meticci tra le due variety sumiominate del 
T. cacao.7' Ciferri's work is a significant contribution to the knowl- 
edge of cacao varieties and their distribution. 
1935: Cheesman describes the branching system and dimorphism 
in cacao, and studies different ways of vegetative propagation. 
1936: Campos Porto publishes information on the species of Theo- 
broma cultivated at the botanical garden of Rio de Janeiro, with a 
photograph of the inflorescences of T. speciosum; the other species 
referred to are T. cacao, T. bicolor} T. grandiflorum, T. subincanunij and 
T. microcarpum. 
1937: Standley lists and describes five species of Theobroma in his 
Flora of Costa Rica plus a Herrania as T. purpureum. He considers 
T. simdarum, T. angustijolivm, T. bicolor, and T. cacao to be wild in the 
forests and T. leiocarpum as probably wild. 
1937: P£rez Arb61aez publishes a manual for the cacao growers of 
Venezuela including a botanical introduction with descriptions of and 
information on Venezuelan cultivars; he also gives interesting historical 
data. 
1938: Pound finds interesting varieties of cacao in his explorations 
in South America, especially in the upper Amazon basin. 
1938: Bondar publishes a documented book on the cultivation of 
cacao in Bahia. He gives comments and a key to the known species 
of Theobroma. He considers that the varieties cultivated in Bahia for 
6SO-695—64 8 
408 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
many years belong to T. leiocarpum; a great uniformity in the prod- 
uct exists, but he believes that recent introductions of Criollo may 
well stop this uniformity. He also describes and illustrates several 
forms of Forasteros known in the region, 
1939: Diels publishes a new species, T. calodesmis, found by Hertha 
Schultze-Rhonhof in the rain forests of eastern Ecuador, which he 
relates to T. speciosum and T. bernouiUiL 
19S9: Cope's investigations in Trinidad on agents of pollination 
using thrips {FranklinieUa parvula Hood), red ants (Wasmannia 
auropunctata Rog.), and apliids {Toxoptera aurantii B. de Fousc.), 
suggests that red ants and thrips are the responsible agents of pollina- 
tion in a cacao population at River Estate. 
1940: E. W. Eimnart publishes the Badianus Manuscript, the 
earliest book ever written on Mexican medicinal plants, the work of 
two Aztec Indians, Martinus de la Cruz who composed the work in 
Aztec, and Juannes Badianus who translated the text into Latin. 
On plate 68, illustrating six plants, plant no. 2 represents "Tlapal- 
cacauatl," colored cacao, i.e., "tlapal" = colored, "cacauatl" = cacao, 
according to Emmart (p. 273), who adds, "This picture is the earliest 
illustration of the cacao, Theobroma cacao L., the source of chocolate," 
This interesting, primitive drawing clearly illustrates the Criollo 
variety. 
1940: Ducke summarizes his experience of many years in Brazilian 
cacaos with a new and detailed key to the Brazilian species. He 
brings new data into consideration in his classification, as for example 
fruit characters that were unknown before. He gives photographic 
illustrations and new information, based on direct field observations, 
about morphological, phonological, and ecological features of the 
species treated, which are T. cacao, speciosum, spruceanum, micro- 
carpum, obovatum, subincanum, and gmndifiorum. He also includes 
in Theobroma the genus Herrania with one species, T. Mariae. Con- 
cerning T. cacao, Ducke recognizes it as indigenous throughout the 
central and western Amazonia. Ducke considers 2\ leiocarpum 
Bernoulli a mere form of T, cacao and makes the new nomenclatural 
combination: T. cacao L. forma leiocarpum. 
1942: Schery publishes a new species, T. asclepiadijlorum, based 
on specimens from Panama. 
1944- Cuatrecasas publishes a new species with yellow flowers, 
T. cirmolinae, found by the author in the rain forests of the western 
slopes of the western Andes in Colombia. 
1944- Cheesman makes a thorough examination of the taxonomic 
situation in cacao, the most important conclusion being that the whole 
assemblage of wild, semiwild, and cultivated cacao constitutes "one 
interbreeding population." He still supports the main division of 
CUATRECASAS—CACAO AND FTS ALLIES 409 
cacao into two groups of varieties, Criollo and Forastero. He pro- 
poses the new theory that the Criollo, which may occur wild in some 
regions from southern Colombia to southern Mexico, may have 
originated at the headwaters of the Amazon. He divides this group 
into Central American and South American Criollos. The Forasteros 
are divided into Amazonian Forasteros, which can be found wild in 
Amazonia and which are widespread in cultivation, and the Trini- 
tarios, possibly originating from the mingling of South American 
Criollo and Amazonian Forastero stocks. Theobroma pentagonum is a 
simple form of T. cacao, probably a segregate of the large cross- 
fertilized population. The same opinion is expressed with regard to 
T. lewcarpum, which, according to Cheesman, does not belong to 
Amelonado; it is an aberrant form of the Criollo, for which rea- 
son the binomial falls into the strict synonymy of T. cacao. The 
data assembled and arguments of Cheesman are a very valuable 
contribution. 
1946: Chevalier publishes a monographic revision of Theobroma. 
He recognizes 13 species (excluding Herrania) arranged according to 
the five sections of Bernoulli. He includes in the first section (Cocoa) 
only T, cacao, in section II, Oreanthes: T. guianensis and T. spruceana; 
in section III, Rhytidocarpus: T. bicolor, T. glauca, and T. Bemouittii; 
in section IV, Telmatocarpus only T. microcarpa and in section V, 
Glossopetalum: T. sylvestris, T. obovata, T. Jerruginea, T. grandijlora, 
T. angustifolia, and T. simiarum. In the key (Tableau Analytique), 
the species are differently arranged; they are divided in the two sec- 
tions of Schumann: Eutkeobroma with four species (T. cacao, T. 
bicolor, T. BemouHlii, and T. glauca), and Bubroma with the other 
nine species. The characters given in the key are not always well 
chosen, and some species are misplaced in the sections (as, e.g., T. 
glauca and Bernouillii). The nomenclature and typification of the 
species are not always correct; the concepts of T. speciosa Willd. ex 
Spreng., T. guianensis (Aubl.) Gmel., T. velutina Benoist, T. syl- 
vestris Mart., T, sylvestris (Aubl.) Don., T. obomta Klotzsch ex 
Bernoulli, T. Jerruginea Bernoulli, T. sinuosum Pav6n and others 
are actually not clarified. Some confusion is also brought with the 
new names T. sagittata Pav6n, T. has fata, and T. undvlata. I have 
identified T. sagittata Pav<5n as Herrania nitida and suppose that T. 
hastata Cheval. is a lapsus calami for the former and T. undvlata 
Cheval. a lapsus calami for T. sinuosum. 
Special treatment is devoted to T. cacao, with which Chevalier 
had long and sound experience. He considers all cultivated cacaos as 
belonging to a single species, T. cacao L., in which four different races 
or "species jordaniennes" can be recognized. These races, which 
cross among themselves l'infini," can be only distinguished by their 
410 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
fruits and seeds. These are lacking in herbaria, and therefore the 
races must be studied in the field. Four of the races or Jordanian 
species of Chevalier correspond to formerly described species (T. 
sativa, T. leiocarpa, T. pentagona, and T. sphaerocarpa) but he adds a 
fifth based only on foliage, T. sagittata, which, as I have pointed out, 
is not a Theobroma. Chevalier's classification for the "formes jorda- 
niennes" of T. cacao L. follows: 
Leaves obovate-oblong, acuminate. 
Fruits ovoid-oblong 5-10 ridged, ± rugose-bullate, long-attenuate into a 
point .  T. sativa 
Fresh seeds yellowish white var. leucosperma 
Fresh seeds dark violet .    var. melanotperma 
Fruits ovoid, rounded at apex, smooth or with 5-10 shallow furrows . T. leiocarpa 
Fruits globose, more or less smooth, rounded or depressed at apex . T. sphaerocarpa 
Fruits ovoid-oblong narrowed to the apex, with 5 very prominent ridges. 
T. pentagona 
Leaves narrowly oblong, or oblong-acuminate, more or less undulate . T. sagittata 
Chevalier goes on to describe the three first "races," and declares that 
he had insufficient information on the other two. Chevalier uses for 
what he calls races, Jordanian species, or Jordanian forms, the same 
binomial denomination as for species, for instance, T. cacao L. forma 
T. sativa which should be T. cacao L. forma cacao according to the 
present rules of nomenclature. It seems right to consider the original 
species described by Linnaeus as belonging to the Criollo form, but 
nothing is clarified by Chevalier using the binomial T. sativa, because 
this name was based on T. cacao L. Chevalier considers his T. sativa 
originally from Central America. To the "Jordanian form" T. 
leiocarpa Bernoulli, Chevalier refers the "Cacao creoulo" of Sao Tomd, 
the Cumacaco, Calabacillo, and Trinitario, and ho supposes it origi- 
nated in Guiana and Brazil. Concerning the Jord an on T. sphaero- 
carpa Chevalier, described on Sao Tom6 (Africa) plants, very similar 
specimens have been found in Venezuela (var. sambito), in the high 
Amazonian forests, and at the Rio Marafi<5n. The Jordanon T. 
pentagona has never been found wild; it seems to be originally from 
Central America. The experience and opinions of Chevalier have 
to be taken into account when considering the classification and origin 
of cultivated cacaos. 
19J$: Cuatrecasas publishes T. capillijerum discovered on the 
Pacific coast of Colombia. 
1947: Llano G6mez publishes information about the cultivated 
cacao in Colombia, with several plates in color representing the 
principal types. 
1948: Rombouts discusses Theobroma SaJUzmaniana Bernoulli, 
showing that it might be based on a flower with defective or abnormal 
CUATRECASAS—CACAO AND ITS ALLIES 411 
petals and therefore cannot be distinguished from other forms of 
T. cacao. 
1949: Standley and Steyermark consider five species of Tkeobroma 
in their Flora of Guatemala, recognizing T. pentagonum and T. 
leiocarpum as different from T. cacao following Bernoulli. T. angusti- 
jolium is given as cultivated and T. bicolor as uncertainly native. 
1949: Cuatrecasas and Le(m describe a new species, T. mammosum, 
collected by Ledn as a rarity on the Atlantic coast of Costa Rica. 
1950: Holdridge publishes some new information on Mexican and 
Central American species of Theobroma, with a key to nine species 
and one Herrania. He suggests that T. pentagonum might be the 
original type and source of the cultivated cacao in Mexico and Central 
America and that the Criollo types were the product of interbreeding 
of T. pentagonum with the South American T. leiocarpum. 
1950-1953: Cuatrecasas publishes T. stipulatum and T. nemorale 
from the rain forests of the Pacific coast of Colombia and T. gileri 
from the Pacific range of Ecuador. 
1951: Freytag publishes a revision of Ouazuma, which helps in the 
study of its relationships of Theobroma. The genus is reduced to 
four species. 
1951: Addison and Miranda Tavares explain the results of their 
six-year work in trying to produce hybrids from different Theobroma 
species. They crossed T. cacao with all the Amazonian species of 
Theobromat without success, and proceeded then to cross the other 
Amazonian species. In 1946, from 719 pollinations of T. speciosum 
on T. cacao, they obtained 29 fruits and 979 seeds, which were mostly 
abnormal and did not germinate. Same results were attained by a 
few pollinations with Herrania mariae. Among 798 cases of polli- 
nation of T. microcarpum on T. cacao 11 fruits and 26 seeds were 
produced, from which only three seedlings were produced which 
grew no more than 10 cm. Similar negative results were produced 
from T. cacao X obovatum and T. bicolor X cacao. Some particular 
trees of T. cacao were more receptive than others; one of them gave 
fruits when submitted to pollination from all other species. When 
T, cacao was used as pollinator on T. microcarpum, T. speciosum, and 
H. mariae, no fruits or seeds were obtained. In 1947, another series 
of cross-pollinations were made on T. cacao with similar results, 
although a few more or less viable hybrids were produced, e.g., 
T. cacao X microcarpum gave 28% fruits, but these decayed after 
developing one month. 
Better results were attained by Addison and Miranda in crossing 
T. grandifiorum and T. obovatum; many hybrid seedlings were pro- 
duced and several developed into perfect trees (in 1% years); the 
leaves, fruits, and flowers of the hybrids showed intermediate char- 
412 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
acters, and the pollen grains were normal and fertile. Well-developed 
hybrids between T. grandiflorum and T. subincanwn, T. obovatum and 
subincanum, and T. speciosum and T. sylvestre (= spruceanum) were 
also produced. In 1948, some fruits were obtained by crossing 
T. cacao with T. grandiflorum, but these gave very few seeds, from 
which only few plants developed up to 15 cm. 
Addison and Miranda also made grafting experiments with good 
results using T. grandiflorum, T. obovatum and T. subincanum. T, 
bicolor, T. speciosum and T. sylvestre (spruceanum) proved to be 
another successful grafting group. 
During their experiments, Addison & Miranda had the opportunity 
of making interesting morphological and physiological observations. 
The seeds of Theobroma usually germinate within 15 days. Tkeobroma 
subincanum, obovatum, grandiflorum, microcarpum, and II. mariae 
were found to have hypogeous germination, whereas T. cacao, sylvestre 
(spruceanum)} bicolor, and speciosum have it epigeous. The very 
young leaves are green in T. speciosum, sylvestre, bicolor, and micro- 
carpum, but they may be green or red in the other species. 
1952-1953: The Anglo-Colombian Cacao-Collecting Expedition pub- 
lishes reports on its explorations in the search for wild and cultivated 
species of Theobrctona and Herrania in Colombia. The expedition took 
place from June 1952 to October 1953, with the participation of the 
British botanists and specialists F. W. Cope, D. J. Taylor, R. E. D. 
Baker, P. C. Holliday, and B. G. Bartley. The Colombian botanists 
who joined the expedition wore H. Garcia Barriga, Canuto Cardona, 
R. Romero Castaneda, and Alvaro Fernandez P. The main areas 
explored were: (1) parts of the rivers Caquetfi, Apaporis, Vaup6s, 
Negro (Guainfa), Imrida, and their tributaries in the provinces of 
Amazonas and Vaup6s, from 1°30' S. to 3° N. and from C7° W. to 
71° W.; (2) parts of the rivers Putumayo, Caquetd, and Cagu&n in the 
provinces of Caquetfi and Putumayo, from 0°20' S, to 2° N. and from 
74° W. to 77° W.; (3) parts of the trans-Andean provinces of Valle del 
Cauca and El Choc<5, from 3° N. to 6° N. and between 76° W. and 78° 
W.; (4) scattered areas in the provinces of Antioquia, Norte de 
Santander, Magdalena, Santander, and ITuila. 
The expedition made 191 botanical collections, of which 63 were of 
living material sent to Trinidad. The well-preserved specimens have 
been extremely useful for the study of the species and their 
geographical distribution. Twelve indigenous species of Theobroma 
were collected (T. calodesmis, microcarpum, subincanum, grandiflorum, 
obovatum, capilliferum, gileri, nemorale, cirmolinaet simiarum, stipu- 
latum, and chocoense). T. bicolor, always found planted, and T. cacao 
were also collected. In a very few areas (Rio Cagu&n, Rio San Miguel) 
spontaneous trees of cacao were found inside the forest but under 
CUATRECASAS—CACAO AND ITS ALLIES 413 
circumstances that make it impossible to say with complete certainty 
that these trees were spontaneous. In general the subspontaneous 
and planted cacaos found in the southeastern region of Colombia 
were of the very uniform Amelonado type. The information and 
materials (living and preserved) gathered by this expedition are a 
very important contribution to the knowledge of Theobroma. 
1954: Ducke makes a revision of his previous synopsis of the 
Brazilian species, incorporating new morphological data into an 
accurate, precise, well-balanced key. He introduces the character of 
the ramification being 5-whorled and 3-whorled in separating T. cacao 
from the other seven Brazilian species (T. bicolor, T. speciosum, 
T. spruceanum, T. microcarpum, T. obovatum, T. subincanum, and 
T. grandijlorum). Ducke considers each fertile stamen as the union 
of two or three (stamens geminous and trigeminous), and as Addison, 
Molina, and Pires had already observed before, characterizes T. 
spruceanum as having geminous stamens. Ducke still retains Herrania 
in Theobroma, with two Brazilian species, T. Mariae and T. Ca- 
margoanum (Schultes) Ducke. He summarizes the ecology and dis- 
tribution of the genus in Brazil, calling it a typical Amazonian genus; 
he writes that it is not absent in any place in Amazonia where rain 
forest exists. 
1956: Cuatrecasas recognizes seven species of Theobroma for the 
Flora of Peru: T. calodesmis, T. grandijlorum (planted), T. obovatum, 
T. speciosum, T. subincanum, T. bicolor, and T. cacao subsp. leiocarpum, 
which is found spontaneous in the rain forests of Peru. 
1958: Schultes publishes the results of his discoveries and research 
on Herrania. His synopsis comprises 17 species, eight of them new. 
One species spreads northwards to Costa Rica, and the others are 
limited to the humid tropics of South America. This monograph 
shows the consistent unity of the group Herrania and the consistency 
of the characters that may be used to separate it from Theobroma and 
other related genera. 
1958: Mora Urpi found a much greater variability in T. cacao 
throughout Mexico and Central America than in South America. In 
Central America and southern Mexico there can be found today prac- 
tically all the known forms of cacao, for which reason Mora believes 
that Central America has been the center of domestication of the 
cultivated cacao; historical data also support this theory. He con- 
siders cacao as having been probably introduced in South America in 
pre-Colombian times. The geographical distribution of the Criollo 
type would also prove this theory. The author agrees with Holdridge 
in considering the pentagonum form as playing an important role in 
the origin of the cultivated hybrid complex; he considers pentagonum 
native in Central America and the original and most ancient form 
414 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
of T. cacao, from which, through mutation, introgressive hybridiza- 
tion, and geographical isolation, the present population arose (p. 34). 
1959: Soria confirms the observations of Mora about the great 
variation in the characteristics of shells and seeds of Theobroma cacao 
in plantations in Nicaragua. Trying to establish the correlation be- 
tween pod shape and color of the seeds, he found that dark-colored 
seeds occurred in a large percentage of Criollo type pods, and white 
seeds were often found in pods of the Forastero and Calabacillo types. 
This agrees with the previous observation by Mora of dark seeds in the 
T. pentagonum pod-type. These observations, according to Soria, 
show that it is probable that the genetic factors controlling these 
characters are independent of each other. Soria sees as reasonable 
Holdridge's theory that Criollo cacao is a result of crosses between T. 
pentagonum and T. Iciocarpum. But he adds "The possibility cannot 
be overlooked, however, that the CriolZos originated as mutations in 
populations on the periphery of the area of distribution of the species, 
the mutations afterwards being fixed and maintained through geo- 
graphic isolation and selection. In this case, pentagona could be a 
product of mutations in Criollo cacaos." Soria emphasizes that 
pentagonum can be fertilized very easily in either direction by other 
types of T. cacao. Mora observed such hybrids as often having con- 
spicuous characteristics of pentagonum. "All these observations lead 
to the conclusion that pentagona is nothing more than one of the ex- 
tremes in the variability of the complex of types forming the species 
T. cacao." 
I960: At the River Estate Experiment Station of the Imperial 
College of Tropical Agriculture in Trinidad, 13 species of Theobroma 
were planted for research and observation. Cope and Bartley suggest 
the possible interrelationships of species of Theobroma, distributing 
them in two groups: 1) with epigeal germination and growth con- 
tinuing from below jorquette, comprising T. cacao with 3-5-branched 
jorquette and T. bicolort speciosum and calodesmis with 3-branched 
jorquette; 2) hypogeal germination and growth continuing from above 
jorquette in T. microcarpum, grandijlorum, subincanum, obovatum, 
angustifolium, mammosum, simiarum, cirmolinae, and nemorale. This 
is the first attempt to classify the genus on the basis of germination 
and branching. 
1960: Crist6bal publishes an excellent monograph on Ayenia with 
important information concerning the relationships with other genera 
of Sterculiaceae. 
1960: Le6n, in Hardy's Cacao Manual, summarizes the taxonomy 
of Theobroma, recognizing 19 species and considering as doubtful T. 
kalagua, tessmannii, ferruginea, and glauca. He gives abridged de- 
scriptions and distribution for T. cacao, bicolor, bernouUlii, capUliferat 
CUATRECASAS—CACAO AND ITS ALLIES 415 
calodesmis, asclepiadijlorum, microcarpa, gileri, guianensis (= speciosa 
sensu Chevalier), spruceana, angustifolia, cirmolinae, grandijiorat 
mammosa, obovata, simiarum, stipvlata, sylvestris (— subincana sensu 
Chevalier), and nemoralis. The fruits of 17 species are illustrated. 
In the classification he follows more or less Chevalier; under T. cacao 
he distinguishes three subspecies: sativa (Lam.), leiocarpa Bern, and 
pentagona (Bern.); the listed cultivars are classified according to van 
Hall. 
1961: Soria reports on cacao in Mexico, having visited extensive 
plantations in Tabasco. Before 1900, the variety cultivated in Mex- 
ico was almost exclusively Criollo, but at present that is disappear- 
ing, hardier and more productive varieties being substituted. He 
observes great variability in the pod form in plantations of old 
Criollo, which always have white seeds. Great variation is also 
seen at present in the widespread hybrid populations that resemble 
the Trinitario of South America, although the Mexican types lack 
the red pigmentation of the shells usually exhibited by Trinitario. 
(They are mostly whitish green or slightly reddish.) 
Morphology 
Stem and branching (Fig. 1).—There is a dimorphism in the 
vegetative organs of Theobroma. The main stem and the adventi- 
tious orthotropic shoots have a radial structure, and the normal, 
plagiotropic branches are monopodial and dorsiventral. The trunk 
is sympodial. 
The seedlings have an erect stem with regular, long-petiolate leaves 
arranged in phyllotaxy cycles of 5/13, 5/8, or 3/8 (Cook, Baker). 
After reaching a height of a few feet the vegetative end of the stem 
stops growing and by the way of a cluster of secondary buds it 
forks into 3 to 5 spreading branches arranged in a terminal whorl 
called a "jorquette" or "fan." These branches are plagio tropic and 
dorsiventral, with alternate, distichous, short-petiolate leaves with 
a phyllotaxy of 1/2. Further growth of the stem may now take place 
by two different ways: 1. One of the dormant buds axillary to the 
branches of the jorquette, and therefore adjacent to the central, 
inert, apex of the stem, develops into a new vertical shoot with the 
same structure as the main stem and looking like its continuation. 
It grows to a limited extent, ending also with a whorl of 3 to 5 branches; 
from above this second whorl or jorquette a third shoot is developed 
in the same way, forming a third internode of the stem, and so on. 
By this way a sympodial main trunk is built, with alternating inter- 
nodes and nodes with regularly centered verticils of branches. Since 
new terminal shoots are produced above the jorquette next to the 
416 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
apex we can call this pseudapical growth. 2. No buds at all develop 
above the first whorl of branches but an adventitious lateral one below 
it grows into a vigorous upright shoot with a structure similar to that of 
the main stem. Although lateral, it forces the node of the jorquette 
to one side, and takes progressively the central position of the stem, of 
which it will appear to be the continuation. After reaching some 
length it forks, ending in a jorquette; a new adventitious shoot is 
formed below that jorquette and so successively a sympodial trunk is 
built, with alternating internodes and irregular nodes. In this case 
the trunk is usually not truly straight and the whorls of branches, in 
spite of the fact that these tend to take a circular position around the 
stem, are always more inclined to one side, often making a lateral 
bunch; the closer to the jorquette the lateral adventitious shoots 
originate, the less irregular is the appearance of the sympodium 
resulting. We can call this sub terminal growth. 
The dimorphism of the stems is transmitted by the buds. Those 
of the seedlings and upright (orthotropic) shoots produce only, again, 
orthotropic shoots (chupons) bearing long-petiolate leaves and pro- 
ducing only the plagiotropic, dorsiventral branches of one terminal 
jorquette. The buds of the lateral, plagiotropic branches produce 
only other plagiotropic branches. Only exceptionally due to special 
physiological conditions or following mechanical injuries (e.g., trim- 
ming), do plagiotropic branches originate upright shoots (chupons). 
More exceptionally the extraordinary formation of alternate plagio- 
tropic branches has been observed on upright stems which have 
failed to form a jorquette (Baker 1961, p. 9), but this has to be con- 
sidered an abnormal case due to unknown special conditions of some 
cultivated trees. 
The lateral, plagiotropic branches are monopodial and branch by 
axillary buds; frequently the growth of a lateral branch bends the 
young joint of the primary branch forcing this into an angle, thus 
simulating a dichotomous fork; branches may appear several times 
forked and are then called ' 'dichotomous'' branches. 
The stem and branching dimorphism is important in the practice 
of propagation and cultivation of Theobroma trees, because only the 
trees produced by cuttings of orthotropic stems (chupons) are upright 
and regular; conversely, those from plagiotropic (dorsiventral) lateral 
branches, branch bilaterally (dorsiventrally) and tend to slant or to 
bow (incline), being thus weaker. In cultivated cocoa the formation 
of adventitious upright shoots (chupons) on branches and at the base 
of the trees is frequently observed; they may be used in practice to 
regenerate old trees by pruning, and as cuttings for propagation. 
But the production of chupons is always too small to serve for ex- 
CUATRECASAS—CACAO AND ITS ALLIES 417 
X 
1 
* 
VI 
N 
? 
hi 
& 
¥ 
Figure 1.—Stem growth in Tktobroma trees, a, b, subterminal or subapical growth; a, 
adult seedling of T. cacao with its primary stem (bearing long-petiolated, radially 
arranged leaves) topped by a whorl of dorsiventral, leafy branches; b, formation of the 
sympodial trunk in T. cacao by way of upright adventitious shoots from buds borne 
below the terminal verticil (or jorquette). c, d, pseudoterminal growth: c, formation 
of the sympodial trunk by developing one of the axillary buds of the terminal whorl 
of branches (e.g., sect. Glossopetalum); d, apex of stem, topped by a whorl of 3 branches 
each with an axillary bud, of which only one will develop (growth above jorquette). 
tensive propagation. The two different kinds of stem growth have 
taxonomic implications. 
418 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Leaves.—There is a dimorphism of leaves correlated with the 
dimorphic stems. The leaves are arranged in several phyllotaxic 
cycles (5/13, 5/8, 3/8 have been recorded) around the radial, ortho- 
tropic stems or shoots, and are distichally alternate (1/2 phyllotaxy) 
on plagiotropic (lateral) branches. 
The first leaves (on orthotropic stems) are long-petiolate and sym- 
metrical. The petiole is elongate and thickened at both ends, forming 
a long, cylindrical pulvinus below the lamina and a more tubercular, 
shorter one at the base; this normal type of petiole facilitates all 
kinds of orientation to the blade. 
The leaves of plagiotropic branches (the normal ones of the mature 
tree) are short-petiolate and asymmetrical. The petiole is with very 
few exceptions reduced to the thickened part of the cushions. The 
blade has more or less markedly unequal halves, especially at the 
base, which may be extremely asymmetrical. 
The blades are simple and pinnatinerved, thick-coriaceous or 
chartaceous, with a strong midrib and several alternate, spreading- 
ascending, prominent secondary nerves; there are elevated tertiary 
nerves. Lesser ones form a small usually conspicuous reticulum. 
Often the lowest pair of secondary nerves is somewhat more separated 
from the next pair than the others, and may give some impression of 
a trinerved base; sometimes there are one or two stronger developed 
tertiary nerves, giving some appearance of a 5-nerved or 7-nerved 
base, but usually the main costa is much stronger than the secondary 
nerves and these are thicker than the tertiary, so that the mainly 
pinnate arrangement is always clear. In some cases, as in the primary 
leaves of T. bicolor, the lateral lower nerves are more clearly arranged 
so as to show a 5-7-nerved base, and some botanists describe them as 
palmatinerved. 
The shape of the blades is often ovate, obovate-elliptic-oblong, 
or lanceolate, and usually acuminate at apex and obtuse or rounded 
at base, but there is a great deal of variation from one species to 
another. The margin is basically entire, but sometimes slightly 
sinuate or broadly dentate in adult leaves; the primary leaves may be 
coarsely dentate in the upper half, Indurnent is present except in 
a few species; most species have a more or less dense tomentum on the 
underside, which may be composed of one, two, or three different kinds 
or sizes of stellate hairs. This tomentum may cover the whole lower 
surface of the leaves entirely or may cover the areoles between the re- 
ticulum leaving all or part of the venation glabrous. The different 
kinds of hairs and their distribution supply good taxonomic characters. 
Inflorescence (Fig. 2).—The inflorescence is of the definite type. 
In some cases it may be a well-branched dichasium as in T. bicolor, but 
generally, the dichasium is totally or partly reduced to a monochasium 
CUATRECASAS—CACAO AND ITS ALLIES 419 
Figure 2.—Inflorescence in Theobroma: A, caulinar inflorescence of T. betnouillii subsp. 
capilliferum (Cuatr. 16160). b, caulinar inflorescence in T. glaucum (Cope & Hoi. 118). 
c, diagrammatic terminal inflorescence branch in T. cacao. d, detail of sympodia. 
branch of inflorescence in T. cacaot diagrammatic, after Stahel. e, inflorescence of 
T. cacao, diagrammatic, after Stahel, f, c, detail of inflorescence in T. cacao, dia- 
grammatic, after Stahel. h, inflorescence of T. bicolor (Klug 2021). i, diagrammatic 
inflorescence of T. bicolor after Stahel. 
420 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
[Figure 3] 
CUATRECASAS—CACAO AND ITS ALLIES 421 
of the cincinnate type. Frequently the branches are very short, 
forming nodose, articulated sympodia, in which the internodes are 
hardly noticeable and the bracts appear almost imbricate. Usually 
the bracts are alternate and the fertile terminal branchlets or peduncles 
end with 3 bracteoles and one pedicel, the 3 bracteoles corresponding 
to a theoretical terminal dichasium which develops only the central 
flower. According to Stahel the peduncles have bilateral, the pedicels 
radial structure. 
The inflorescences may be axillary, in young branchlets, but more 
often are originated on short, woody branchlets on the trunk and 
branches; these perennial branchlets form irregular tubercles, some- 
times very protuberant, which may form woody branchlets up to 
several centimeters in length, producing flowering cymes at their 
ends. The flowers are always pedicellate, the pedicels being relatively 
long, longer than the reduced branchlets of the cymes. 
Calyx (Figs, 3, 4).—The five sepals are valvate and may be almost 
free and spreading at anthesis or united from one fourth to one half 
or more of their length; the lower united part is cupular, the free parts 
are patulous-reflexed at anthesis but finally the whole calyx becomes 
reflexed, exposing the inner surface. In some instances, the sepals 
unite 2 by 2 simulating a calyx of 3 lobes, two of them twice as broad 
as the third. In the section Andropetalum the sepals usually are united 
by three and two and together form a two-lobate cup. The calyx 
is persistent and its remains may often be seen below premature 
fruits. 
In most cases the sepals are tomentose outside with abundant stel- 
late, ochraceous or ferrugineous hairs, but they may also be puberulous 
or glabrous, as in species or forms of the sections Telmdtocarpus and 
Theobroma. In the latter multicellular, glandular, stipitate trichomes 
are present. The upper or inner surface of sepals is often glabrous 
or may be more or less pubescent. The inner margin always has a 
Figure 3-—Calyx and aestivation in Theobroma: a, b, calyx of grandifiofum (Cuatr, 
25801) with sepals united by pairs appearing to be trimerous. c, calyx of 7\ cacao 
(Cuatr. 26004), the sepals spreading, very shortly united at base. D, e> calyx of T♦ 
nemoraU with semispreading and semireflexed sepals, united more than r, c, calyx 
of 7\ chocoense9 with reflexed sepals unequally united in their length; the basal 
glandular pap ilia s very conspicuous. H, diagrammatic long, section of bud in T. 
citmolinae showing the relative position of flower parts in section Glossopetalum; at 
the left side the folded petal (the alternating staminode cut away), at right the staminode 
(the alternating petal cut away), i, globular bud of T. ckocoenst with valvate aestiva- 
tion, pedicel, bracteoles and peduncle apparent (X 2). j, ovoid, elongated bud of 
7\ velutinum (X 2). k, petals in bud showing the contorted aestivation in T. velutinum 
(Benoist 161)t X 5, l, diagrammatic long, section of bud in 7\ cacao showing the rela- 
tive position of flower parts in sections Ottantkesy Tkcobroma, and Rhytidocarpus. 
422 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Figure 4*—a-c, three types of floral diagrams in Thtobroma: a, sections Glossopttdum 
Andropetalum and Oreantkes (part); s, sections Thtobroma, Rkytidocarpus and Oreamhes 
(part); c, section Telmatocatpus. D, basic diagrammatic representation of vascular 
bundles in the flower of the Byttnerineae; the marginal vessels (bundles) derive from the 
vascular branches directed to the petals; the fertile stamens are epipetal, the sta ml nodes 
episepal; p> vascularization of petal in continuous line, after Gazet du Chateller slightly 
modified, e, vascular bundles in flower of Theobroma (sect. Glossopetalum); bundles 
5n stamens branching shortly above the base; adapted from Gazet du Chatelier, p, 
section of an ovary in Theobroma (X 20). G, gynoecium in T, cirmolinae (X 5). H, 
flower in anthesis with spreading sepals in T. syfoeitre (Ducke 7882), X 2, i, flower in 
CUATRECASAS—CACAO AND ITS ALLIES 423 
narrow band of extremely minute and dense, whitish, stellate hairs 
which join the sepals before an thesis. At the base of the sepals, 
inside, there are commonly glandular, stalked trichomes, which may 
be very scarce but often are numerous and dense, forming a ring 
outside the place of petal insertion. These glandular trichomes are 
lacking in some species or very rare and scattered above on the sepals. 
Corolla (Figs. 3, 4).—The 5 petals are free and uniform; their 
special feature is that they are strangulated into two very different 
parts united by a narrow joint. The lower part is cymbiform and 
erect, rather carnose and rigid, usually 3-7-(or 1)-nerved and has the 
appearance of a hood rounded at the top; in fact, it represents the 
claw of the petal, and because of its appearance, it is called the 
"hood" {cucvllus). The upper part of the petal (the lamina) is flat, 
varying in shape from oblong to elliptical or discoid, membranaceous 
or very thick, yellow, red or purplish. It is almost sessile and directly 
articulate to the apex of the hood or may be supported by a narrowly 
laminar pedicel, which is its basal extension; in some species the lamina 
is lacking or is almost reduced to the pedicellar extension (T. mam- 
mosum). The petals are dextrorsely contorted in estivation, the 
laminae being erect when directly articulated to the hood or horizontal 
and reflexed through the folding of the pedicel. In an thesis the 
laminae are erect. 
Androecium (Figs. 3, 4).—This is formed by two verticils, which 
are connate in a tube at the base. The sterile outer whorl has 5 
petaloid staminodes, which are subulate, oblong, or obovate, and 
usually very showy with the same color of the corresponding petal- 
laminae. They may be erect or reflexed in estivation, and erect, 
spreading like a star, or reflexed in an thesis; they are thick-membra- 
naceous or carnose and firm, or when subulate or narrowly lanceolate 
they may have a thick, carnose midrib; they may be glabrous, but 
more commonly are hairy or covered by minute, nmricate trichomes. 
The inner whorl consists of 5 fertile stamens, with thick filaments 
which are connate to the tube except for a short (1-3 mm.) free part; 
the apex is 2- or 3-furcate and each short branch bears an anther. The 
filaments are spreading and curved and, being opposite the petals, 
each anther is concealed in the cavity of its corresponding opposite 
petal-hood. The anthers are 2-celled, and each cell, ellipsoid or almost 
globose, is unilocular and opens by a longitudinal slit. 
The 2- or 3-antheriferous stamens of Theobroma have been treated by 
some botanists as a result of the coalescence of two or three original 
anthesis with reflexed sepals, spreading petals and erect staminodes in T. cirmolinae 
(Cuatr. 15336), X 2. j, flower of T. cirmolinae (Cuatr. 15336) initiating anthesis in 
apical view (X 2). 
680-685—64 i 
424 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
[Figure 5] 
CUATRECASAS—CACAO AND ITS ALLIES 425 
stamens. Nevertheless, I consider them to be bifurcate or trifurcate 
original stamens. The anatomical works of Gazet du Chatelier (1940, 
p. 278) prove this assertion; at the base of the flower, 5 vascular 
bundles proceed to the five staminal filaments, these bundles being 
forked above to serve the two short branches of the filament in 
T. cacao. 
Gynoecium (Figs. 3, 4).—This is of the coeno-syncarpic type, 
superior, with carpels opposite to petals. The ovary is 5-celled with 
axile placentation and many ovules in two rows in each cavity; it is 
ovoid or ellipsoid, more or less markedly 5-ridged and furrowed, 
densely stellate-tomentose, or rarely glabrous or covered by stipitate 
glands. 
Sty lodes 5, free or more or less adherent to one another, simulating 
a single style, glabrous, usually about twice as long as the ovary, thin 
and ending in a punctiform stigmatic apex. The ovules are anatropous 
with dorsal raphe and two integuments. 
Fruit and seed (Figs. 5-7).—The fruit is almost baccate or sub- 
drupaceous and indehiscent, the various types differing in the firmness 
of the pericarp and in the shape. Almost always there can be dis- 
tinguished three layers in the pericarp. In the sections Glossopetalum 
and Andropetalum, the fruits are externally rigid, hard, the epicarp 
being woody, about 1 to 2 mm. thick, with an outer tomentose epiderm; 
the mesocarp is fleshy, differing little in color and firmness from the 
adjacent endocarp; the inner surface of the latter is a thin but com- 
pact membrane; sometimes, the whole endocarp is reduced to this 
membrane. When the fruit is ripe the carnose inner layers decay or 
dry, and shrink, but the rigidity of the epicarp maintains absolutely 
the size and shape of the fruit, keeping the loose seeds inside if they 
have not been accidentally liberated. In the section Ekyiidocarpus, 
the mesocarp is the rigid, woody layer; the epicarp being thinner and 
carnose, although also with an outer tomentose epiderm; the endocarp 
is also carnose, and also provided with an inner membrane. In the 
section Oreanthes, typified by T. speciosum, the whole pericarp is 
5 to 6 mm. thick; the innermost layer, the endocarp, although very thin, 
Figure 5.—Fruit sections of Tkeobroma species: a, long, section of fruit of T, sitntarum 
(Cuatr. 26515A), X %• b, transection of pericarp of a, natural size, c, transection of 
pericarp of T. grandiflorum, X 1. D, transection of pericarp of T. bicolor, X 1. e, 
transection of pericarp of T, gileri, X 2. f, transection of pericarp of T. speciosum, 
X 1. <5, transection of pericarp in T. cacao cultivar "cundiamor" (Cuatr. 26492), X 1. 
H, transection of pericarp in T. cacao fma. pentagonum (Cuatr. 26540), X 1. i, transec- 
tion of pericarp in T. cacao cultivar '' la gar to" (Cuatr. 26004), X 1. J, transection of 
pericarp in Herrania cuatrecasana (Cuatr. 25793), X 2. ed, epiderm; ep} epicarp; mt 
mesocarp; en, endocarp; pi, interior pelicule limiting the endocarp inside; pu, pulp; 
s, seed. 
426 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
V 
i1; ^ 
itoiMv jk /'V I 
■h -.v 
,r 
u 
\ 
J 
t 
I 
H 
i 
\ 
\ ^ 
4V 
a 
* 
v 
* 
c 
Figure 6.—Seeds of Theobroma, natural size: a-e> T, grandiflorum (Cuatr. 25780T): a, 
front view of seed stripped from pulp; b, c, d, side and front view of embryo; E, seed 
surrounded by its pulpy layer, f-h, T* simiarum (Cuatr, 26515A): f, seed; g, embryo; 
H, embryo tn apical view, i, j, T* mammcsum (Cuatn 25791): seed in lateral and apical 
CUATRECASAS—CACAO AND ITS ALLIES 427 
is woody and rigid; the mesocarp is thick and fleshy; the epicarp is also 
woody but thin and less compact. When the mesocarp dries or decays, 
the epicarp shrinks slightly, becoming more or less rugose. In the 
section Telmatocarpus the innermost layer, the endocarp, is the most 
compact and rigid, although thin; the epicarp is coriaceous and thin 
and the mesocarp is thick and fleshy with strong bundles which build 
protruding ridges and veins covered by the epicarp. Finally, in 
section Theobroma (Cacao) the fruit is almost baccate because the 
whole pericarp is carnose; the inner membrane and the outer epiderm 
may be very firm but the whole pericarp decays easily; it can also dry 
out, being then coriaceous. Usually the three layers are conspicuous, 
the epicarp carnose and thick, with glabrous outer epiderm, the 
vascular mesocarp papyraceous, rigid, thinly woody, and the endocarp 
carnose, more or less thick, with an inner pellicle; in some forms of 
cacao the mesocarp may be reduced to a very thin or discontinuous 
layer or to isolated vascular bundles (fig. 5i); rarely the endocarp is 
reduced to the inner pellicle (fig. 5h, forma pentagonum). Usually, 
the dorsal vascular bundles of each carpel develop into transverse 
membranes within the fleshy epicarp connecting with the mesocarpial 
layer. Gummy sacs are always present in several parts of the pericarp. 
The young fruit, as well as the ovary, has five cavities with the in- 
cipient seeds arranged in one or two rows in each cavity. At maturity 
the cell walls vanish and the seeds with their thick outer pulpy layer 
fill the single cavity, arranged usually in five rows. 
The shape and size of the fruit are variable and they are, com- 
bined, specific characters except for cultivated cacaos. The fruits 
range between 6 and 35 cm. long by 5 to 12 cm. broad and may 
contain, when ripe, from 16 to 60 seeds. They may be ellipsoid, 
globose, ovoid, oblong, or fusiform, with rounded or attenuate ends, 
with completely smooth surface like a potato or marked more or 
less with 5 or 10 ridges, or they may be echinate or verrucose. In 
all cases they are indehiscent and the liberation of seeds follows the 
decay of the shell, which in many cases, as in those with hard, woody 
pericarp, may take so long that the seeds have died. The common 
natural way of propagation of the seeds is accidental, usually by 
view, k-m, T. angustifolium (Cuatr. 25790): k, seed stripped from pulpy layer; l, 
embryo; m, embryo in apical view, n-p, T. gileri (Cuatr. 26167): n, seed in apical 
view; o, same laterally; p, embryo, q-s, T. speciosum: q, seed; r, embryo; s, embryo 
in transection, showing the folding of cotyledons, t-v, T. bernouUlii subsp. capilliferum 
(Cuatr. 17034): t, seed; u, embryo; v, embryo in apical view, w-xx, 7*. cacao fma. 
pentagonum (Cuatr. 26004): w, seed; x, embryo; xx, transection of embryo, v, yy, 7*. 
cacao fma. pentagonum (Cuatr. 26540): Y, transection of seed with episperm and pulp; 
yy, seed, z, zz, T. cacao cultivar "cundiamor" (Cuatr. 26492): z, seed; zz, transection 
of seed with episperm and pulp. 
428 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
animals (mostly monkeys, but also squirrels, rats, and other animals) 
which break the pericarp in order to suck the pulp surrounding the 
seeds, which may be expelled later in other places, thus disseminating 
the seeds (fig. 5). 
The fruits, which are commonly called pods in English, mazorcas 
in Spanish and cabosses in French, may stay on the tree or fall down 
after maturation; in the latter case, they may fall with the peduncle 
(e.g., T. bicolor) or without it (as in T. grandiflorum). Precise obser- 
vations in many species are wanted. 
The seeds are ellipsoid, ovoid or amygdaloid, more or less irregularly 
compressed, complanate, or terete, and range from 15 to 40 mm, long 
and 10 to 22 mm. broad; the integuments or skin form two strata 
with an additional outer, thick, gelatinous-pulpy layer surrounding 
them. The testa is generally thick and subcoriaceous, with an 
external epiderm covered by a thick cuticle, a thick layer of poly- 
hedric and mucilaginous cells, and an inner layer of sclerosed cells. 
The inner tegument is a thin membrane of several layers of thin- 
walled, complanate cells. Inside, the large embryo is composed of 
two large, thick, strongly folded and corrugate cotyledons and a 
straight, rather thick, terete radicle, the plumule being scarcely 
developed. The endosperm at maturity has the shape of a very fine 
pellicle, containing scattered cells with calcium oxalate, covering the 
embryo outside and between its foldings. The cotyledons possess 
an epiderm, often with scattered, stipitate glandular trichomes and 
a main cellular tissue rich in starch, fat, aleurone, tannoid and alka- 
loid al substances, among these the important theobromine compounds. 
In most species the cotyledons are white, but in a few they are violet, 
reddish, purplish, being stained by tannins (figs. 5, 6). Germina- 
tion may be either epigeous or hypogeous according to the species 
(fig. ?). 
The pulp surrounding and united to the seeds is white, yellowish, 
or yellow, and often sweet and aromatic and palatable, but it may 
be also scentless and tasteless to men; it is, however, always appreci- 
ated by animals, which hunt the pods, extract the seeds to suck the 
pulp, thus disseminating them. 
In appropriate conditions the pulp suffers a fermentation process 
which separates it from the seed; during that fermentation, very well 
known in the case of T. cacao, chemical changes take place inside the 
embryo developing a special aroma. In the industry of cacao torre- 
faction completes the desired effects of fermentation. 
Premature fruits keep their viability for some time provided they 
are protected against loss of humidity and stored under suitable 
temperatures (20-25° C.); when ripe, the seeds become immediately 
ready for germination; they may germinate inside the pods. The 
CTJATRECASAS—CACAO AND ITS ALLIES 429 
Figure 7.—a, Epigeal germination in Tkeobroma bemouillii subsp. capilliferum (Cuatr. 
17350A). B, Hypogeal germination in T. grandiflorum (Cuatr., Cope & Bart. 25780A). 
germinating power of Tkeobroma seeds lasts only for a short time, a 
few weeks; observations on T. cacao have shown a maximum extension 
time of viability to about three months, when carefully preserved in 
their pods or under special protection. They are extremely sensitive 
to the degree of humidity, which has to be kept high, and to low 
temperatures- Recent experiments in Turrialba showed that cacao 
seeds could not resist low temperatures for even a short time; seeds 
exposed for 16 minutes at 8° C. lowered the germinating capacity to 
6%, and 4 minutes exposure at 2° C. inhibited germination almost 
completely (Hunter, 1959; Hunter & Boroughs, 1961). This may 
be the explanation why cacao seeds lose their germinating power 
430 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
when transported by airplanes at high altitude (Hunter, personal 
communication). 
Ecology 
Theobroma is a tropical American genus restricted to the lower 
tree-story of the evergreen rain forests. The species demand a high 
degree of mean annual temperature with narrow oscillations, a con- 
stant high humidity, and protection (shade) against direct radiation 
and evaporation. Several species are often found at the edges of 
rivers or marshes in more or less temporarily flooded areas; others 
always grow on elevated, drained places. They like relatively rich 
and neutral soils. These conditions are found only in the warm, wet, 
forested, equatorial regions between latitudes 18° North and 15° 
South, with temperatures of 20° to 30° C., with a minimum of 16° and 
maximum of 40°. A few species grow at higher altitudes up to 1250 
meters, being able to withstand minimum temperatures of 14° and 
even 12° C. Where Theobroma is at home, the rainfall is from 2000 
mm. to 8000 mm, annually or even higher, and is more or less evenly 
distributed throughout the year. Theobroma does not resist even 
short dry seasons without the protection of dense shade and local 
humidity. In cultivation T, cacao can endure less humid climates, 
and more open lighter spots especially when irrigated, and somewhat 
lower temperatures than the normal optimum. So the area of culti- 
vated cacaos may extend far above 20° North and below 20° South 
of the Equator. Not only the Theobroma, trees but also the seeds are 
highly specialized to the humid equatorial ecological conditions. It 
is known that the seeds keep for a very short time their capacity for 
germination, which often takes place inside of the pod; only under 
high humidity and optimum temperatures can they maintain their 
viability (see above). 
Geographic Distribution 
The genus Theobroma is a typical neotropical genus, distributed 
throughout the rain forests of the western hemisphere between 
latitudes 18° North and 15° South. Some species have a broad range 
of distribution, like T. subincanum, which is spread throughout the 
Amazon-Orinoco basins, being one of the most ancient of the genus. 
The elevation of the Andes in the early Tertiary separated populations 
of Theobroma previously widespread before, favoring speciation 
through isolation. Vicarian species separated by this way are T, 
subincanum (east of the Andes) and T. hylaeum (west); T. microcarpum 
(east) and T. gileri (west). The complexity of the mountains of the 
northern part of Colombia through Central America was also an iso- 
lating factor which favored speciation in that part of the hemi- 
sphere where regional or local endemics are present. Maps 1 and 2 
are self-explanatory. 
CUATRECASAS—CACAO AND ITS ALLIES 431 
Map 1.—Geographical areas of Theobroma sect. Oreantfus: 2, T. sylvestre; 3, T. speciosum; 
4, T. telutinum; 5, T. glaucum; T. bernouiilii: 6a, subsp, bernouillii; 6b, subsp. as- 
clepiadifloTum; 6c, subsp. capilliferum. 
t T^7 
sac 
aT,& 
vfv 
up: 
A.' % » 
A 
t 
* 
* 
t 
22 K+ 4 
14 
I3| 
19] 16 
201 
15 ?- 
> 
\ 
Map 2.—Geographical areas of Theobroma sect. Glossopetalum (10-21) and sect. Andro- 
petalum (22). 10, T. angustifolium; 11,7*. cirmolinae; 12, T. jtipulatum; 13, T. choco- 
ffwf; 14, 7*. jt-midfUfft; 15, T. grandifioruni; 16, 7*. obovatum; 17, 7". subincattutn; 18, 
T. hylaeum; 19, T. nsmorale; 20, T, sinuosum; 21, 7*. canumanense; 22, T. m-ammosum. 
432 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Pollination 
Although transportation of pollen by the wind has had some ac- 
ceptance, it seems that only pollination by insects has been proved. 
Works by Harland, Stahel, Posnette, Saunders, Cope, Uzel, Jones, 
and Pound, have demonstrated that several kinds of flying and crawl- 
ing insects are involved in pollen transportation, among them thrips, 
ants, midges, and aphids. Experiments by Cope proved that Frank- 
liniella parvuta Hood and Wasmannia auropunctata Rog. were mostly 
responsible for pollen transportation in Trinidad. Saunders found in 
Costa Rica that Forcipomyia midges performed pollination in cacao. 
The aphid Toxoplera aurantii Fouse is also recorded as a pollinating 
agent. 
Relationships 
The basic features of Theobroma are: Flowers bisexual, pentamer- 
ous; sepals valvate; petals strangulated, contorted in bud with 
cymbiform, cucullate lower half; 5 stamens opposite to^petals*and 5 
evident, alternating staminodes united in a short basal tube; stamens 
shortly 2-3-branched; anthers 2-celled; ovary superior, 5-celled; 
ovules many, anatropous with 2 integuments; fruit subdrupaceous or 
subbaccate; seeds with pulpy envelope; cotyledons folded, corrugate; 
evergreen trees with dimorphic branching and dimorphic, entire, 
alternate leaves. This diagnosis places the genus in the family 
Sterculiaceae (fig. 8). 
Theobroma exhibits a unique set of characters which makes it a 
very "natural" genus. However, some of the outstanding features of 
its floral structure are also shared by other genera, the most con- 
spicuous being the cucullate or concave lower part of the petals which 
define the tribe Byttnerieae DC., and determine the close relation- 
ships between its members: Byttneria, Ayenia, Rulingia, Commer- 
sonia, Theobroma, Guazuma, Herrania, Abroma, Glossostemon, Scapho- 
petalum, and Lepionychia. In most of these cases the similarity with 
Theobroma in the flower structure (petals, androecium-tube, stami- 
nodia, and position of anthers) is so obvious that it was noticed since 
early times. The first historical association of Theobroma to another 
genus was by Linnaeus who joined it with Guamma under Poly- 
adelphia Pentandria. Lamarck (1785) was the first to make a family 
associating Theobroma with Abroma, Guasuma, Ayenia, Byttneria, and 
Kleinhovia. Jussieu (1789), associated Theobroma with Abroma, 
Guasuma, Byttneria, Dombeya, Assonia, Pentapetes, and Melhania in 
sectio V [bis] of ordo XIV. Kunth (1823) was the first to estab- 
lish critically the main groups of the Sterculiaceae, one of them the 
Byttneriaceae verae including Theobroma, Guasuma, Abroma, Glosso- 
CUATRECASAS—CACAO AND ITS ALLIES 433 
stemon, Byttneria, Ayenia, and Commersonia, This grouping was 
basically followed by DC. (1824), Endlicher (1840), although he 
separated the Sterculiaceae from Byttneriaceae, Baillon (1873) in his 
series Byttneri^es, Bentham & Hooker (1862), who enlarged the 
family to 7 tribes, and Schumann (1890) who enlarged it to 8 tribes. 
The latter botanist, who made an outstanding contribution to the 
comparative morphology and taxonomy of the whole family, did not 
alter the concept of the Byttnerieae DC. as presented by Bentham and 
Hooker. Recent workers, like Gazet du Chatelier (1940), who made 
broad comparative anatomical and morphological studies in the 
Sterculiaceae, found good reasons to keep Schumann's basic taxonomic 
approach. 
Bentham and Hooker divided the tribe artificially in two groups 
which were named by Schumann Theobrominae and Byttnerinae, with 
respectively 2-3-antheriferous and 1-antheriferous stamens. On the 
other hand, the four genera of Byttnerinae differ from Theobroma also 
because Byttneria has spirally convolute cotyledons, short, dentiform 
staminodes and linear, rather thick, petal-laminae; Ayenia has very 
long, linear petal-claws, trilocular anthers, and spirally convolute 
cotyledons; the Old World Commersonia and Rulingia have a pitcher- 
shaped petal base and flat cotyledons. From the other Theobrominae 
genera, Theobroma is distinguished by the special structure of the 
petals, staminodia, and vegetative system; the Persian genus Glos- 
sostemon is a shrub with hairy, dentate leaves, ovate-oblong petals, 
concave at base, and with many short stamens connate to the basal 
part of the staminodes; the Old World Leptonyckia differs by its short, 
squamiform petals, fertile stamens with filaments much longer than 
the staminodes, and flat cotyledons; the west African shrub Scapho- 
petalum has exappendiculate petal hoods, a campanulate androecium 
with shortly triangular staminodes and sessile 3-grouped anthers; the 
tropical American Guazuma differs, besides in the fruit, by the long, 
bifid petal appendages, the spirally convolute cotyledons, and the 
vegetative structure, the leaves being serrate; Abroma, an oriental 
genus spread from eastern India through the Pacific islands to Aus- 
tralia, is similar to Theobroma in the floral arrangement but usually 
has more developed petal laminae, shorter petaloid staminodes, 
subsessile anther groups, flat cotyledons, a different vegetative habit, 
and usually cordate, more or less lobate, hairy leaves. Moreover, 
all genera mentioned of the Byttnerinae differ from Theobroma by 
their capsular, generally dehiscent fruit. Only Theobroma and 
Herrania in the tribe have an indehiscent baccate or subdrupaceous 
fruit. For this reason, Schumann united them, calling the latter 
section Herrania, an arrangement adopted by other botanists, such 
as Pittier and Ducke. Nevertheless, Bernoulli, the monographer who 
434 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
[Ficure 8] 
CUATRECASAS—CACAO AND ITS ALLIES 435 
went deeply into the genus, and Chevalier in his revision consider 
Herrania and Theobroma different genera. R. E. Schultes followed 
the same line in monographing, after long experience, the genus 
Herrania with 17 species. They are undoubtedly two well-defined, 
independent genera. 
Herrania} Abroma, Guazuma, and Byitneria surely are the genera 
closest to Theobroma. Chromosome number and palynology help to 
determine relationships. The chromosome number is identical for 
Theobroma and Herrania, 2n = 20 but it is 2n — 16 for Guazuma and 
2n ~ 14 in Byitneria (Crist6bal); data for Abroma not available. The 
pollen grains are suboblate in Theobroma, prolate in Herrania, prolate- 
spherical in Cruazuma, and oblate in Abroma. 
Because of the similarity of the fruits and the confusion which had 
prevailed in the past between Theobroma and Herrania, their differ- 
ences are summarized here as follows: 
Theobroma. Stem sympodial, with 3-5-verticillate branching; 
branches dimorphic; branching copious; leaves dimorphic; leaf-blades 
simple, entire; petal lamina more or less rounded to lanceolate, not 
more than twice as long as the hood, erect or inflexed and contorted 
in aestivation; pollen grains suboblate; cotyledons strongly folded and 
corrugate; fruit usually smooth or rugose, angular, seldom strongly 
costate; staminal filament symmetrically and shortly 2- or 3-furcate 
at apex. 
Herrania. Stem monopodial, unbranched, with apical growth; 
leaves uniform, 5-9 digitate, long-petiolate, in a terminal, lax cluster; 
petal lamina many times longer than the hood, linear, pendulous 
in anthesis, involute in aestivation; pollen grains prolate; cotyledons 
thick, flat or very slightly folded; fruit usually strongly costate; 
staminal filament usually asymmetrically parted in two branches, 
one 1-antheriferous, the other 2-antheriferous (fig. 9). 
Evolution 
The question of how the genus Theobroma may have originated 
is & speculative matter on which botanists like Schumann (1886) 
Figure 8.—Genera related to Theobroma: a-h, Guazuma: a, articulated and hooded petal 
with bifid appendix, from inside; b, same from outside; c, same in lateral view, X 5; 
d, androecium, X 5; is, fertile stamen, X 20; f, sepals, X 2; g, bud; h, gynoecium, X 5 
(G. tomeniosa, Cuatr. 22942), i, j, Byttneria: i, articulated petal from inside, X 10; 
j, same laterally, k, flower, X 10. l, gynoecium, X 10. u, androecium, X 10. 
N, detail of the anther, X 20. o, sepals, from inside and outside, X 5. p, bud, X 5. 
q, carpel of fruit from inside, outside and apical view, X 2 (B. arguta, Cuatr. 8226). 
r-2, Abroma: r, articulated, hooded petal, X 2; s, same in lateral view; t, hood from 
inside and u, from outside, X 5; v, sepal, X 2; w, pistil, X 5; x, androecium surround- 
ing the gynoecium, X 5; y, base of staminode with laterally attached stamens, X 5; 
z, biantheriferous stamen, X 10 (Abroma augusta, Sulit 18880). 
436 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
[Figure 9] 
CUATRECASAS—CACAO AND ITS ALLIES 437 
did not want to take a stand. Edlin (1935) who developed a theory 
on the evolution of the Mai vales, considers the family Sterculiaceae 
limited to the tribe Sterculieae, all other groups forming the family 
Byttneriaceae; he considers the stamens the result of union. Gazet 
du Chatelier (1940), after a detailed examination of the Sterculiaceae, 
came to the conclusion that there was an original unknown type from 
which were derived two diverging groups (subfamilies or families), 
the "Eriolaenees" and the "Buettneriees," but he did not go much 
further in his speculative evolution; the stamens of Theobroma are 
considered as branched by him. 
All the genera of the By ttnerieae are similar and probably originated 
at the same time evolving from an original unknown type; they 
diversified their flowers and the leaves, probably through mutations 
aided by geographical and ecological barriers. The parts have 
evolved independently, e.g., Rvlingia has undivided fertile stamens, 
a more ancient character, but pitcherlike petals, a more evolved one. 
Conversely, Leptonychia has simple, more primitive petals but 
exhibits branched stamens, a more advanced character. Byttneria 
and Guazuma have elaborate, advanced petals but less developed 
staminodes; Commersonia, as well as Abroma and Theobroma, have 
more advanced petals and staminodes than the other genera. 
ScaphopetaluTn is an example of a more advanced type due to the loss 
of the petal lamina and reduction of the staminodes. Even if we can 
attribute primitiveness or the contrary to some characters, it is not 
possible to draw a lineal series of genera according to antiquity. 
Nevertheless, I would venture to say that Theobroma and Herrania 
belong to the most modern in the By ttnerieae because of the structure 
of the fruit, with thick and partly or totally carnose pericarp and 
delicate, short-lived seeds. These may be characters acquired in 
the process of evolution and kept by their adaptation to the extremely 
hot and humid ecological conditions of the tropical American forests. 
It also seems to me that Herrania is a more evolved genus in regard 
to the flower, but not in the simplicity of the monopodialf juvenile- 
Figure 9,—a-h, Herrania fulcherrima v. pacifica (Patino 23): a, articulated and hooded 
petal, X 5; B, segment of androecium with a staminode and the adjacent stamens, 
X 5; c, bud, X 5; D, stamens, X 10; e, gynoecium, X 5; F, seed, X 1; g, transection of 
seed, X 1; h, embryo, X 1. i-k, H. cuatrecasana: i, embryo; j, seed; k, transection 
of embryo, X 1- l-r, Theobroma bicolor (Garcia B, 11178): L, petal from inside, 
outside and laterally, X 5; m, androecium, X S;N, fertile stamen and part of stamina! 
tube, X 10; o, sepal from inside and outside, X 2; p, gynoecium, X 5; q, styles, X 5; 
r, bud and pedicel, X 2. a-Y, T. syhestre (Ducke 7882): s, petal from inside and 
laterally, X 5; T, androecium, X S; u, stamen, X 10; v, bud, X 2; w, gynoecium, X 5; 
y, sepal from outside and inside, X 2. 
438 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
like, unbranched stems, in the digitate, loosely clustered leaves, and 
inseparable, hypogeous cotyledons, due probably to the stringent 
ecological conditions of the shadowy underlayer of the humid tropical 
forest. Aside from Herrania, the sections of Theobroma (Glossopetalum) 
with pseudoterminal growth, may be more primitive than the ones 
(Oreanthes, Rhytidocarpus, Theobroma) which have lost the axillary 
buds of the jorquette branches, necessitating lateral shoots to continue 
growing. The section Theobroma may be more evolved than the 
others on account of the 5-branching system and carnose fruits. The 
section Telmatocarpus may be more advanced in another direction 
because of the reduction or absence of the petal lamina and the 
discontinuity of the vascular, woody system in the pericarp, which is 
only partially woody and more vulnerable. The parallelism in evolu- 
tion of the sections Theobroma and Telmatocarpvs is seen in the 
glabrous or almost glabrous leaves, more suited to rain-forest ecology. 
The section Rhytidocarpus may be an ancient type with less showy 
petals and staminodes, axillary flowers, and a thick-woody pericarp. 
No fossils belonging to Theobroma have been recorded. 
The geographic distribution does not give any solution to these 
questions of evolution because almost all sections are represented at 
both sides of the Andes. It seems that the richest region in species 
is around Panama and Colombia, where species with a very restricted 
area are found, especially if we consider this region extended to 
Costa Rica. I feel that Theobroma is a genus with a marvelous set 
of characters controlled perhaps by independent genes, which seem- 
ingly can combine independently resulting in many different sets of 
combinations. 
Economic Uses 
The seeds of Theobroma are rich in starch (15%), protein (15%), 
and oil (50%), for which reason they are considered a substantial food. 
Moreover, they have a volatile oil (cacao-essence) which gives an 
aromatic flavor and 1.5 to 3% of theobromine, an alkaloid known for 
its stimulant properties. Caffeine is also present in Theobroma seeds. 
Both alkaloids have been found in the seeds and leaves of T. bicolor, 
cacao, grandiflorum, microcarpum, obovatum, speciosum, sylvestre, and 
subincanum (Willaman and Schubert, 1961). The cacao seeds contain 
also a red pigment, tanine, and small quantities of malic and tartaric 
acids, asparagine, and coline. 
It is not necessary to emphasize the economic importance of the 
industry in cocoa and chocolate. Most of the wild species of cacao 
are often used by the natives, who suck the pulp or prepare refreshing 
drinks with the pulp. The seeds of most species may serve for the 
preparation of chocolate, but actually only one species has become 
CTJATRECASAS—CACAO AND ITS ALLIES 439 
commercially important in this respect, T. cacao, which is the only one 
widely cultivated. An important secondary product from cocoa 
seeds is the cocoa butter extracted by pressure during the process of 
making chocolate. Cocoa butter is important in cosmetics and 
pharmaceutical industries. Cacao extracts and theobromine are 
important in medicine because of their cardiotonic and diuretic 
properties. 
The wood of several Theobroma species is important in local con- 
struction and because of its toughness and strength is very much used 
in the manufacture of tools and parts of instruments and machines. 
Anatomy of the Wood 
Contributed bt William L. Stern 1 
This study of the wood of Theobroma is based largely on microscope 
slides borrowed from the S. J. Record Memorial Collection of woods 
at Yale University and from the wood collection of the Imperial 
Forestry Institute at the University of Oxford in England.2 It is 
regrettable that among these slides, only 9 species were present 
(table 1). However, the description of the wood probably represents 
a fairly good outline of at least the qualitative aspects of the micro- 
scopic structure, and is sufficiently complete to enable comparisons 
between Theobroma and other genera to be made. 
It is evident from this brief study that noticeable variation occurs 
in the wood anatomy of different specimens of the same species. In 
this regard it is interesting to note that Record and Hess (1943, p. 
517) were impressed with the structural variation in rays in different 
parts of the same specimen in their study of the woods of Sterculiaceae. 
As a whole, however, the wood of Theobroma species does not present 
any characters of significant anatomical import which would enable 
us to separate them on anatomical grounds, Chattaway (1937) also 
found this to be true of the genera she studied in her investigation of 
the Sterculiaceae (sensu Edlin 1935). 
> Rwehincks: 
Bailey, L W. The problem of differentiating trachelds, fiber-trachelds, and llbrlfarm wood fibers. 
Trop. Woods 54:15-23. 1936. 
Bentimm, G., 6 Hooker, J. D. Sterculiaceae, In Genera plantarnm. 1:214-228. London, 1862. 
Chattaway, M. Margaret. The wood of the Sterculiaceae. L Specialisation of the vertical wood 
parenchyma within the sub-family Sterculleae. New Phytol, 31:119-132, 1932. 
 , Ray development In the Sterculiaceae. Forestry 7:93-108. 1933. 
 , The wood anatomy of the family Sterculiaceae. Phil. Trans. Royal Soc. London. Ser. B-Blol. 
Sd. 228:313-366. 1937. 
Edlin, H. L. A critical revision of certain taxonomlc groups of Malvales. New Phytol. 34: 1-20. 
1935. 
Metcalfe, 0. R., 6 OhsJt, L. Anatomy of the dicotyledons, 1:261. Oxford, 1960. 
Record, S. J„ & Hess, R. W. Timbers of the New World, p. £17. New Haven, 1943. 
> I would like to thank Dr. Qraeme Berlyn of Yale and Dr. L. Chalk of Oxford for theirklndnessln making 
slides available for study. 
680-6955—64 5 
440 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
The imperforate tracheary elements are all fiber-tracheids; that is, 
the bordered pits are smaller in diameter than those found in the walls 
of vessel elements in the same species. Bordered pits usually show 
extended inner apertures; these may be crossed or not. The wall 
thickness varies from thick to thin, sometimes even within the same 
species (cf* specimens of T. tricolor). 
Pores are distributed mainly in the solitary configuration on the 
transverse section (34-86 percent; average 62 percent); radial mul- 
tiples are next in abundance (8-47 percent; average 33 percent) and 
pore clusters are least abundant (0-10 percent; average 4.5 percent). 
In different specimens of the same species, dissimilarities may occur; 
for example, in T. obovatum {Williams 161), solitary pores account for 
86 percent of the pores per field, whereas in T. obovatum (Williams 230), 
they account for only 57 percent of the pores per field. Perforation 
plates are entirely simple. Vessel element end walls form angles from 
45° to 80° with the vertical. Intervascular pitting is alternate. The 
Table 1.—Specimens examined in anatomical analyses 
Species of Theobroma Collector and No. Origin j Herb, 
voucher 
USw 
No.* 
Yw 
No. 
FHOw 
NO. 
angustifoUum 
bernouillii Pitt. 
bicolor H. & B. 
tricolor H. & B. 
bicolor H. & B- 
bicolor H. & B. 
cacao Ij. 
cacao L. 
grandiflorum 
(Willd.) 
So hum. 
microcarpum 
Mart. 
obovatum 
Ktotzsch ex 
Bernoulli 
obovatum 
Klotzsch ex 
Bernoulli 
sylvestre Mart. 
subincanum 
Mart. 
subincanum 
Mart. 
Cooper Slater 
242 
Pittier 4105 
For. Dept. Br. 
Hond. 
II. 2192/29(?) 
"Ford-Brazil 
397" (?) 
LI. Williams 
2149 
U. Williams 
3346 
"L. 3225 (via 
Hamburg)" 
Vigne 2433 
LI. Williams 
2401 
Krukoff 6203 
LI. Williams 
161 
LI. Williams 
230- 
Ducke 103 
LI. Williams 
LI. Williams 
Panama 
Panama 
British 
Hon- 
duras 
Brazil 
Peru 
Peru 
South 
America 
Ghana 
Peru 
Brazil 
Peru 
Peru 
Brazil 
Peru 
Peru 
Y 
TJS 
F 
F 
F 
F 
Kumasi: 
K(?) 
F 
US 
F 
F 
Y 
F 
F 
30 
10595 
22075 
17804 
18176 
17893 
36510 
71232 
17263 
21362 
17578 
18144 
3502 
5632 
6998 
7008 
7007 
5703 
6898 
7001 
7010 
7011 
7009 
7000 
6999 
* Abbreviations from W, L. Stern & K\ L* Chambers. The citation of wood specimens and herbarium 
vouchers in anatomical research* Taxon* 0: 7-13. I960. 
CUATRECASAS—CACAO AND ITS ALLIES 441 
bordered pits are frequently crowded and their outlines markedly 
angular. Other times the pits are rounded to elliptical. Vessel-axiaJ 
parenchyma pitting and vessel-ray parenchyma pitting generally 
follow the pattern of the intervascular pitting. Occasionally pits may 
be elongated or slightly irregular. No deposits or tyloses appeared 
in any vessels. 
Both uniseriate and multiseriate vascular rays occur in each of 
the specimens examined. Multiseriate rays may be up to 20 cells 
wide (in T. sylvestre), but a width of 10 to 15 cells is more common. 
Uniseriate rays are much lower in height than multiseriate rays; the 
latter range from 30 to 230 cells high. There is often evidence of dis- 
sociation of these broad, high rays into lower, narrower rays by the 
"intrusive action" of fiber-tracheids while the cells are still in a plastic 
stage. Many of the multiseriate rays are characterized by the 
presence of sheath cells, e.g., in T. obovatum (Williams 161); however, 
they never form complete sheaths about the rays and are rare in some 
specimens. Multiseriate rays are heterocellular, with the multi- 
seriate portion comprising procumbent cells, and uniseriate, alate 
extensions of 1 to several (15+) upright or square cells; uniseriate 
rays are usually homo cellular composed of square or upright cells 
and sometimes occasional procumbent elements. The ray cells are 
commonly characterized by deposits of reddish or yellowish, non- 
staining materials. 
Axial parenchyma occurs in two dispositions: apotracheal, as 
diffuse and/or diffuse-in-aggregates arrangements, and paratracheal, 
as vasicentric sheaths 1 or 2 cells wide. Sometimes a ladderlike 
configuration is formed on the transverse section by short bands of 
axial parenchyma which frequently intercept vascular rays (e.g., in 
T. angustijolium). 
Storying of tissues occurs in the wood of Theobroma, but in its 
most highly developed state, it would have to be considered incon- 
spicuous. Where it does appear, it is limited in distribution and 
confined to the uniseriate rays. In T. microcarpum it was also 
observed in the axial parenchyma. 
Crystals occur in the wood of all species examined. Generally 
they are more frequent and conspicuous in the cells of rays, although 
they also occur in axial parenchyma cells of some species. In rays, 
only crystals of rhomboidal nature were observed except in T. mic.ro- 
carpum where large druses were present exclusively. Crystals in 
axial parenchyma cells are mostly rhomboidal, but in some species 
small druses occur. 
Discussion.—The most significant anatomical studies of the wood 
of Sterculiaceae are those of Chattaway (1932, 1933, 1937). Unfor- 
tunately, her work is of limited value as a basis for comparison here, 
442 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
for she adopted the restricted view of the family proposed by Edlin 
(1935) and confined herself to the tribe Sterculieae. It should be 
mentioned that Edlin suggested dividing the Sterculiaceae, as treated 
by Bentham and Hooker (1862), into two families: Sterculiaceae, to 
be restricted to the tribe Sterculieae, and Bucttneriaceae, to contain 
all other taxa (including Theobroma). Chattaway corroborated 
Edlin's proposals according to the anatomical findings which resulted 
from her studies. Nevertheless, it does not seem to me, judging 
solely from her summary of the characteristic anatomical features of 
the Sterculiaceae (sensw stricto), that the wood anatomy of Theobroma 
(which would be eliminated from Sterculiaceae according to Edlin's 
concept) would preclude its being allied with the species upon which 
she reported if we used only anatomical bases. There are only two 
apparent anatomical differences between Theobroma wood and that 
of Sterculiaceae (sensu stricto): Regardless of statements to the 
contrary (Metcalfe and Chalk 1950, p. 251), the imperforate elements 
in Theobroma wood are not libriform wood fibers, but fiber-tracheids 
with small bordered pits {sensu Bailey 1936). Chattaway describes 
corresponding cells in Sterculiaceae (sensu stricto) as libriform wood 
fibers. Also, she indicates that the rays in Theobroma woods lack 
sheath cells (Chattaway 1932, 1937), "but are present in the rays of 
all genera of the Sterculiaceae except Heriliera." I cannot agree 
that Theobroma rays are totally devoid of these specialized elements. 
Although they are of sporadic occurrence, it is relatively easy to 
demonstrate them among the rays in any given tangential section. 
In summary we can say that Theobroma woods are characterized 
by fiber-tracheids with small bordered pits, mostly solitary pores, 
simple perforation plates, alternate intervascular pitting, both homo- 
cellular uniseriate rays and heterocellular multiseriate rays with 
sheath cells in the same species, both apotrachea! and paratracheal 
axial parenchyma in the same species, and crystalliferous deposits 
which are most abundant in the cells of ray tissue. Storied structure 
is present to a limited degree and is confined largely to the uniseriate 
rays. Although anatomy is variable within specimens of a given 
species, it is not consistently variable to allow for the division of the 
genus on anatomical grounds. In my opinion, the wood anatomy of 
Theobroma does not differ significantly from that in Edlin's Stercu- 
liaceae as delineated by Chattaway. 
Pollen Morphology of Theobroma and Related Genera 
Contributed »t G. Erdtman * 
Theobroma L. (fig. 10): Pollen grains 3-colporate, peritreme, 
suboblate (about 15-22X17.5-25 p). 
* Falynologieal Laboratory of the Swedish Natural Science Research Council, Stockholm. 
CUATRECASAS—CACAO AND ITS ALLIES 443 
Species investigated: T. angmtifolium Moc. & Sess6 (Pittier s.n.): 
about ?X23.5 ju; T. bernouUlii Pitt, (Pittier 4105): about ?X22 p; 
T, bicolor Humb. & Bonpl. {King 2021): about 16X18 p; T. cacao L. 
(CaMeron 107): about 15X17.5 n; T. glaucum Karst. (Holliday & 
Cope T-118): about 22X24.5 p; T. grandiflorum Schum. {Archer 
7549): about 19X22.5 T. microcarpum Mart. {Archer 7551): about 
16.5X21 ja; T. speciosum Willd. var. coriaceum Huber {Rusby 647): 
about 22X24.5 /i. 
r ♦ 
Figure 10,—Palynograms, X 1500: a, Herrartia pulckerrima v. pacifiea Schult.; b, Glos- 
sostemon bruguieri DC,; c, Guazuma polybotrya Cav.; Theobroma glaucum Karst.; 
B, Abtoma augusta L,; G, Erdtman & A. L. Nilsson, original. 
444 CONTRIBUTIONS PROM THE NATIONAL HERBARIUM 
Examples: T. glaucum Karst. (HoUiday c£ Cope X—1 IS): pollen 
grains 3-colporate, peritreme (amb circular), oblate spheroidal (about 
22X24.5 n). Apocolpium diameter about 18 Colpi narrow, about 
18 ft long. Ora about 3.5 ju broad. Exine about 1.5 p thick. Sexine 
as thick as nexine or slightly thicker, tectate. Tegillum distinctly 
undulating. The waves of the tegillum are smoother than those of, 
for example, Herrania pulcherrima var. padjica, but nevertheless they 
impart to the pollen surface a reticuloid pattern with muroid ridges 
(supported by one or two rows of endosexinous bacula) separated by 
luminoid depressions (diameter up to 3 /*)• The tegillar bottom of the 
latter seems to be supported by stray baculoid rods. 
T. microcarpum Mart. (Archer 7551): pollen grains 3-colporate 
(amb circular), suboblate (about 16.5X21 n). Apocolpium diameter 
about 13.5 jtx. Colpi narrow, about 10 fi long. Ora lalongate (about 
1.5X3 ft), Exine about 1.6 ti thick. Sexine thicker than nexine, 
tectate, undulating (waves not as smooth as in T. glaucum). Retic- 
uloid pattern much as in T. glaucum, with more or less irregular 
luminoid areas (longest axis up to about 3.5 %). 
T. speciosum Willd. var. corvaceum Huber (Busby 647): pollen 
grains 3-colporate, oblate spheroidal (about 22X24,5 /*). A single 
4-colporate (loxocolpate) pollen grain seen. 
Apocolpium diameter about 15 m- Colpi narrow, about 12 n long. 
Ora lalongate (about 3X8 /*)• 
Exine about 1.5 ft thick or a little less. Sexine thicker than nexine, 
probably tectate, presenting a reticuloid pattern (OL) with narrow 
straight muroid and irregularly polygonal luminoid areas (maximum 
diameter of the latter 1.5 ft). Muroid areas supported by a single 
row of endosexinous bacula. 
The pollen grains of Herrania differ from those of Theobroma. 
Herrania Goud.: pollen grains 3-colporate, peritreme, prolate 
(about 32-35X23-25 p). 
Species investigated: H. camargoana Schult. (Baker 39): about 
34X25 n; H. cuairecasana Garcia B. (Cuatrecasas 11168): about 
35X24 fi; H. mariae Schum. (Ducke 595 and Martins 318 (type)): 
about 33 X25 n; H. pulcherrima var. padjica Schult. (Patino 23): about 
32X23 p. 
Example: H. pulcherrima var. padjica Schult.: pollen grains 3- 
colporate, peritreme, prolate (about 32 X 23 ju). 
Apocolpium diameter about 14 /i. Colpi about 25 ju. Ora about 
2.25 n high, slightly lalongate, their horizontal margins incrassate. 
Exine about 2 p thick at poles, 1 n at center of mesocolpia. Sexine 
thicker than nexine, tectate. Tegillum undulating, with anastomos- 
CUATRECASAS—CACAO AND ITS ALLIES 445 
ing, slightly winding, crestlike and slightly carinate folds imparting 
a distinct reticuloid LO-pattern to the exine surface. Crests about 
1 ft broad at the poles, gradually more narrow (about 0.5 p or less) in 
mesocolpia. They are supported by a single row of endosexinous 
bacula except at the poles, where there are several rows. The lumi- 
noid, concave areas between the folds of the tegillum are equally 
supported by small endosexinous bacula or baculoid rods (largest and 
longest at the poles). The longest diameter of these areas varies 
between 2 and 5 ft or more. 
The pollen grains in Glossostemon bruguieri are somewhat similar 
to those in Herrania. 
Glossostemon bruguieri DC. (Iraq, Falluja, Haines s.n.): pollen 
grains 3-colporate, peritreme, subprolate (35X28 *»)• 
Apocolpium diameter about 8 n. Colpi about 25 p, constricted at 
the equator, ends rounded, margins thickened. Ora lalongate (about 
8X1.5 f»). 
Exine about 2.1 ix thick at the polos, about 1 p at the equator. 
Sexine thicker than nexine, tectate. Tegillum strongly undulating, 
forming distinct, anastomosing muroid ridges (about 0.5 p wide) 
separated by luminoid areas. In the apocolpia and towards the 
colpi margins the latter are very small (diameter usually not exceeding 
0.5 n); in the mesocolpia they are larger (longest diameter up to 4 /x). 
The muroid ridges are supported by a single or double row of endo- 
sexinous bacula. The tegillar bottom of the luminoid areas is also 
supported by small bacula. Bacula in apocolpia considerably longer 
than those in mesocolpia. 
The pollen grains in Abroma and Guazuma are somewhat similar 
to those in Theobroma. 
Examples: Abroma augusta L. (Assam; herb. Riksmus., Stockholm, 
marked "no, 370"): pollen grains cf. 3-colporate, peritreme, oblate 
(21X29 ti). 
Apocolpium diameter about 23 p. Colpi about 5.5X2.5 n, their 
margins incrassate. Ora not very distinct. 
Exine about 1 fx thick, tectate, very slightly undulating, presenting 
a reticuloid pattern. Muroid ridges low, about 1 m wide, supported 
by a double row of endosexinous bacula and enclosing small rounded 
luminoid areas 1-2 m wide. Under each of the latter is one or several 
endosexinous bacula, 
Guazuma polybothra Cav. (Cuba, Boldo s.n.; herb. Madrid, marked 
"no. 94"): pollen grains 3-colporate, peritreme, prolate spheroidal 
(18.5X16.5 jtt). 
Apocolpium diameter about 12 Colpi about 12 ju long. 
Exine about 1 ju thick, tectate, undulating, with distinct narrow, 
446 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
carinate muroid folds separated by luininoid, concave areas (diameter 
less than 1 /i in apocolpia as well as in mesocolpia). 
Guazuma ulmifolia Lam. (Mexico, Pringle 2570): pollen grains 
3-colporate, peritreme, spheroidal (16/x). 
Apocolpium diameter about 5 ju. Colpi about 14 ju long, 1 n wide. 
Ora lalongate, about 1 n high and 3.5 n wide. 
Exine about 1 /x thick (of the same thickness in apocolpia as in meso- 
colpia, probably tectate (tegillum undulating, exhibiting narrow, 
muroid ridges separated by luminoid areas less than 1 n in diameter). 
Cytology * 
CONTEIBUTED BY F. W. COPE 
Chromosome numbers in Theobroma species.—The first pub- 
lished count of 2?i=20 for T. cacao, the now accepted figure, was 
made by Davie (1933) from studies of mitosis in root-tips. He 
noted that "the chromosomes are very small, quite different from 
Malvaceous chromosomes. A few show median constrictions. Two 
pairs of satellited chromosomes were seen." In 1935, Davie con- 
firmed the diploid number of 20 from studies of meiosis in pollen 
mother cells of T. cacao. 
Confirmation of this number has been made for T. cacao by Carletto 
(1946), Munoz Ortega (1948), Simmonds (1954) and Cope (unpub- 
lished). The first three authors have also shown twenty to be the 
diploid number in other Theobroma species. Carletto counted 20 
chromosomes in T. "leiocarpa," T. speciosum, and T. granaiflorum 
and Munoz Ortega in T. "leiocarpaT. "pentagona," T. bicolor, 
T. microcarpum, T. speciosum, T. simiarum, T. capUliferum, T. 
grandijlorum, T. obovatum, T. angusiifolium and T. drmolinae. Sim- 
monds confirmed 2n~20 in T. bicolor and T. angustifolium. Accord- 
ing to Munoz Ortega, the chromosomes throughout the genus show 
medial, submedial, and terminal centromeres. The chromosomes are 
uniformly small, with size gradations within each species examined. 
The largest chromosomes of T. cacao are 2 p in length; the smallest of 
T. microcarpum only 0.5 /i long. 
* References: 
Carletto, G. M. (1946). O name to do cromosomiosem cacauciros. Bol. Tee. lust. Cacau Bahfa No. 
0, 35-30. 
Davie, J♦ H. (1033), Cytological studies in the Malvaceae a&d related families. Jonrn. Genet. 28:334)7, 
Davie, J. H, (1035). Chromosome studies in the Malvaceae and certain related families II, Genetica, 
17: 487-408. 
MuAoz Ortega, J. M. (1048). Estudios cromosomicos en el gGnero Theobroma L. MSS in library of the 
Instltuto Interamericano de Ciencias Agrlcolas, Turrialba, Costa Rica. 
Slmmotids, N. W. (1054). Chromosome behavior in some tropical plants. Heredity, 8:130-146. 
CUATRECASAS—CACAO AND ITS ALLIES 447 
Pollen Incompatibility 5 
Contributed by F. W. Cope 
The incidence of self- and cross-incompatibility, and self- and 
cross-compatibility in T. cacao was first discovered by Pound (1932) 
when he showed that some trees in Trinidad could not set fruit with 
their own pollen nor with one another's. These self- and cross-incom- 
patible trees needed pollen from a self-compatible tree in order to 
set fruit. Posnette (1945) discovered cross-compatibility between 
self-incompatible types in his studies on cacao trees introduced from 
the upper Amazon into Trinidad. The existence of self-incompatible 
and self-compatible cacao trees has now been established in nearly all 
areas where the species is wild or cultivated. 
Cope has shown, in a series of publications, that unlike most other 
plant species showing incompatibility the site of the incompatibility 
reaction in cacao is in the embryo-sac, and not in the stigma or in 
the style. Pollen tubes in incompatible pollinations grow as fast 
as those in compatible pollinations and deliver their male gametes 
into the embryo-sacs in perfectly normal fashion. It is only when the 
male gametes come to lie in contact with their female counterparts that 
any abnormality appears (fig. 11). According to the genotype of the 
tree or trees involved in an incompatible pollination, either one quarter, 
one half, or all encounters between male and female gametes result in 
failure of the fusion process. When an incompatibly pollinated cacao 
flower falls from the tree 25%, 50%, or 100% of the ovules in the 
ovary show nonfusion; in the first two cases the other fertilized 
ovules in the same ovary show normal fusion between the gametes to 
give a zygote and a triploid primary endosperm nucleus in each. 
The genetic system controlling the nonfusion and fusion of gametes 
in the embryo-sac of T. cacao is now known. Three complementary 
loci appear to be involved, which have been called A, B, and S. The 
first two show simple dominance and recessivity; the S locus carries 
multiple alleles between which dominance and independence relation- 
ships exist. The action of the S locus was first postulated by Knight 
and Rogers (1955), based on results obtained from wholly self-incom- 
patible material. The need for other loci, to act in a complementary 
• References: 
Cope, F, W. (1939). Studies In the mechanism of self-Incompatibility In cacao I. 8th Ann. Rep, on 
Cacao Rea. (1930), Trinidad, 20,21. 
——(1940). Studies in the mechanism of self-Incompatibility In cacao II. 9th Aon. Rep. on Cacao 
Bes. (1939), Trinidad, 10-23. 
 (1068). Incompatibility in Theobroma cacao. Nature, London, 181, 270. 
 (1959). Incompatibility in Theobroma cacao. A Hep. on Cacao Res., 1857-68,7-17. 
"Knight, R., and Bogota, H. H. (1955). Incompatibility in Theobroma cacao. Heredity, 9: 09-77. 
Posnette, A. 7. (1946). Incompatibility In Amaron cacao. Trop, Agriculture, Trln., 22:184-187. 
Pound, F. J. (1932). Studies in fraitfulness in cacao. U—Evidence for partial sterility. 1st Ann. Rep. 
on Cacao Res. (1931), Trinidad, 24,25. 
448 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
■ 
tOinu> 
IQjnv 
Figure 11.—a, Camera lucida drawing of an embryo sac of Theobrotna cacao, fixed 24 hours 
after pollination: one male gamete (cP) is in contact with the egg nucleus (9) and 
the second is moving towards the polar nuclei, p; the darkly-staining synergid cell (sy) 
has been penetrated by the pollen tube; and starch grains (s) are abundant. B, camera 
lucida drawing of an incompatibility fertilized embryo sac of T. cacao, 72 hours after 
pollination: one male nucleus lies in contact with the egg nucleus (top L.H.); and the 
second male nucleus (cf a) is associated, unfused, with one polar nucleus (pj), the other 
polar (pi) having moved away, c, as in b, except that the second male nucleus (cTi) 
and the two polar nuclei (p, and pj) are all dissociated (c?i is the first male gamete 
lying in contact with the egg nucleus), d, as in b, except that the second male nucleus 
is separated from the two coherent polar nuclei. F.W. Cope, original. 
CUATRECASAS—CACAO AND ITS ALLIES 449 
manner with the S locus, was pointed out by Cope (1958) in order to 
explain the emergence of self-incompatible progeny from a cross be- 
tween two true-breeding self-compatible parents. 
The A and B loci both act before meiosis. When both are at least 
heterozygous for the dominant all el e, it is believed that a general pre- 
cursor substance is produced and on this the S locus acts to produce 
very highly specific incompatibility reactions between gametes carry- 
ing the same S allele. The S locus acts both before and after meiosis, 
the premeiotic action giving the overall sporophytic control of incom- 
patibility and the postmeiotic action leading to a gametophytic reac- 
tion between gametes. 
If one or more of the A, B, and S loci become homo zy go us for an 
inactive allele the self-incompatible condition is lost; the tree is then 
self-compatible and cross-compatible with any other cacao genotype. 
A few examples of genotypes of the two classes of tree are: 
Self-incompatible Self-compatible 
A ABBS*., aaBBS,., aaBBS«.t 
AaBbS*.y aabbSi.i 
AABbSx., AABBSf.f 
AaBBSt.f aaBbSf.f 
where S, and S, are two active S alleles and S( is an inactive amorph 
of the S series. 
Self-incompatible genotypes are also cross-incompatible if they 
have common dominant S alleles; if one has alleles independent in 
action and one of these is duplicated in the other as a dominant; or 
if both genotypes, carrying only alleles of independent action, have one 
allele in common. 
Genus Theobroma 8 L. 
Theobroma L. [Gen. PI. 351- 1737; Hort. Cliff. 379. 1737] Bp. PI. 782. 1753; 
Gen. PI. ed. 5, 340. 1754; Benth. A Hook. (1862) 225; Bernoulli (1869) 4; 
Schumann (1890) 86 pro parte; Schumann (1896) pro parU; Chevalier (1946) 
269. 
Cacao [Town. Inst. 660, (. 444' 1700]. Miller, Gard. Diet. Abr. ed. 4th. 1754. 
Tribroma Cook, Journ. Washington Acad. Sci. 5:288. 1915. 
Type.—Theobroma cacao L. 
Flowers hermaphroditic, pentamerous, pentacyclic, diplostemonous. 
Buds globose, ovoid or oblong-ovoid. Sepals 5, valvate in aestivation, 
almost free and spreading or more or less united in the lower part, 
cupular, or united by pairs into one single and two double lobes, or 
rarely in two lobes. Petals 5, dextrorsely contorted in aestivation, 
each one strangulated in two halves: 1) a lower part corresponding 
# TTuobroma is a neuter name and the genus must be neuter by tbe present International Code of Nomen- 
clature. The feminine endings for the species used by @ome authors (De Candolle, Bernoulli, Chevalier, 
etc.) are corrected in this revision into the neuter form except for the original bibliographic references. 
450 CONTRIBUTION'S FROM THE NATIONAL HERBARIUM 
to the claw, rigid and strongly veined with the shape of a hood (cu- 
cullus); 2) an upper part, a flat blade (lamina), articulated to the 
indexed apex of the claw. Androecium in two verticils of five, 
united in a tube at base: an outer whorl with 5 sterile, petaloid or 
linear staminodes, opposite to the sepals; an inner whorl with 5 fertile 
stamens opposite to the petals, the filaments short, minutely 2-3- 
branched, each branch with an anther. Anthers hidden inside the 
petal-hoods, bilobate (bithecate), the thecae unilocular and dehiscent 
by longitudinal clefts. Pollen grains 3-colporate, peritreme, sub- 
oblate (about 15-22 x 17.5-25 fi). Gynoecium 5-carpellar, syncarpic, 
superior, the carpels opposite to petals, the ovary ovoid, pentagonal, 
5-celled with axile placentation, the many ovules in two rows in each 
cell. Stylodes 5, connivent, free or more or less united, filiform. 
Stigmas apical, short, acute. Ovules anatropous with two integu- 
ments and dorsal raphe. 
Fruit large, subbaccate or subdrupaceous, in dehiscent, ovoid, 
ellipsoid or oblong, obtuse or acute, smooth or ridged, rugose or 
tuberculate, the pericarp fleshy or hard and partly woody or coriace- 
ous, the vascular axis thin and vanishing; seeds usually in five rows, 
each one surrounded by a thick, fibrose, pulpy tissue filling the cavity 
at maturity, ovoid, ellipsoid, or amygdaloid, the episperm double, 
thick, subcoriaceous, the outer layer with a trichomatic and gelati- 
nous epiderm developing into a thick, pulpy envelope; embryo straight, 
the radicle cylindrical, inferior; cotyledons thick, strongly plicate- 
corrugate; endosperm usually reduced to a filmy membrane covering 
the cotyledons. Germination epigeous or hypogeous. 
Evergreen tree with the apical growth of the stem limited to the 
production of a terminal whorl of 3-5 spreading branches; sympodial 
growth of the stem attained by adventitious upright sub terminal 
shoots or by pseudoapical shoots from buds axillary to the apical 
branching whorl. Primary branching of stem 3- or 5-verticillate, the 
further branching alternate. Leaves simple, entire, penninerved, 
persistent, coriaceous, long-petiolate and varied in phyllotaxy on the 
primary stems, short-petiolate and distichous on the branches. 
Inflorescences dichasial or monochasial (cincinate), axillary or on 
reduced tuberculiform branchlets on trunk and larger branches. 
Peduncles bracteate, articulate to pedicels. 
Pluricellular trichomes in all species, usually as stellate hairs, 
rarely simple. Globose, stipitate glands present in some species. 
Chromosome number: 2ra=20. 
Suhgcneric classification 
The division of the genus Theobroma in five sections by Bernoulli 
is the best to date. He used the characters of the petal-lamina 
(sessile, stipitate, or lacking), shape of staminodes and their position 
CTJATRECASAS—CACAO AND ITS ALLIES 451 
in the bud, and the number of anthers. A sound combination of these 
characters gives five very natural groups. Schumann's (1886, 1890) 
separation of two sections, Theobroma and Bubroma, according to their 
2-antheriferous or 3-antheriferous stamens, leads to an unnatural 
grouping because the number of anthers for each stamen may vary 
in the same section and even in the same species (e.g., T. glaucum). 
For this reason, the combination of Schumann's with Bernoulli's 
classification made by Pittier (1930) was erroneous, because the section 
Oreanthes cannot be placed in either of Schumann's two groups. 
Chevalier (1946) used both classifications but without trying to in- 
tegrate them. Ducke (1954), who published the best elaborated key 
for 7 Brazilian species, did not pay attention to sections, but he used 
the 3- or 5-whorled branching as a new character to distinguish the 
species. Another character, the epigeous or hypogeous germination 
of the seeds correlated with growth-habit of the tree, was used by 
Addison and Tavares (1951) for distinguishing species, and by Cope 
and Bartley (1960) in classifying them. 
I have applied to the classification the mode of germination, and 
the growth and the branching system for all the species, and I have 
found the sections founded by Bernoulli to be very much reinforced 
by the addition of these vegetative features, and other floral and fruit 
characters unknown before. Epigeous germination and sub terminal 
growth apply to all species of the sections Rhytidocarpus, Ore<mthes, 
and Theobroma, whereas the other sections exhibit hypogeous germi- 
nation and pseudoterminal growing. These vegetative characters 
prove to be very important and basic, being uniform for each section, 
but like other characters, even though constant within the section, 
they are not sufficient to give taxonomic recognition to the two groups 
separated by those characters. The Bernoulli sections are all of 
similar rank, independent and probably of parallel origin. Only the 
new section Andropetalum, based in its extraordinarily broad and 
reflexed staminodes, relatively reduced petal-lamina, and gamosepal- 
ous calyx, seems to be closer to Qlossopetalum than the other sections 
to each other. The fruit structure, as explained above, is also impor- 
tant in the present classification. The following key will give a clear 
idea of the characters of each section. Another artificial key is 
added to facilitate the identification of specimens lacking complete 
information. 
The position of the inflorescences (cauline or axillary), the color, 
size, and shape of petal-lamina and staminodes, number of anthers, 
shape of sepals and indument, form and size of fruits, the outline, 
venation, thickness, and firmness of leaves, and especially the kind of 
indumentum they bear, as well as the pilosity on different parts of the 
flowers, inflorescences, and branchlets, and the form and caducousness 
of stipules are the characters used to distinguish the species. 
452 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Key to Sections of Theobroma 
1. Cotyledons epigeous at germination; growth of stem by adventitious upright, 
lateral-eubterminal shoots; staminodes in aestivation erect. 
2. Staminodes thick-linear, obtuse; petal-hood 1-nerved; petal-lamina sub- 
sessile; stamens 2-antheriferous; pericarp thick, ridged and nerved, the 
mesocarp very hard, woody; primary branches ternate; leaves tomentose 
beneath 1. Rhytidocarpus 
(Contains the single species: Tr bicolor) 
2. Staminodes linear-subulate or lanceolate, acute; petal-hood 3-nerved. 
3. Fetal-lamina sessile; stamens 3- or 2-antheriferous; pericarp coriaceous; 
primary branches ternate; leaves tomentose beneath . . 2. Oreanthes 
3. Petal-lamina attenuate-stipitate; stamens 2-antheriferous; pericarp firmly 
carnose; primary branches quinate; leaves glabrous or puberulous 
beneath  3, Theobroma 
(Contains the single species: T. cacao) 
1. Cotyledons hypogeous at germination; growth of stem pseudoapical; primary 
branches ternate; stamens 3-antheriferous. 
4. Staminodes flexuose in bud, ovate, subulate-caudate; petal-hood 5-nerved; 
petal-lamina lacking; pericarp carnose-coriaceous, lignose-ridged and 
reticulate; leaves glabrous or puberulous 4. Telraatocarpus 
4. Staminodes reflexed in bud, obovate-oblong, broadly lanceolate or broadly 
obovate, reflexed or erect in anthesis; petal-hood 7-nerved; pericarp 
rigid, the epicarp hard, woody; leaves tomentose beneath. 
5. Staminodes oblong-obovate or lanceolate, reflexed or erect at anthesis; 
petal-lamina broadly developed, flat, stipitate; sepals more or less 
united and reflexed 5. Glossopetalum 
5. Staminodes broadly obovate, as broad as long; petal-lamina somewhat 
reduced, narrow, and plicate; calyx cupular, the sepals united one 
half or one third their length 6. Andropetalum 
(Contains the single species: T. mammosum) 
KEY TO SECTION OREANTHES 
I. Leaves stellate-tomentose beneath on the minor reticulate veins, the areoles 
densely, minutely, stellulate-tomentose. 
2. Filaments 2-antheriferous; inflorescences small, on the leafy branches; flow- 
ers rather small (sepals 7-9 x 2-2.5 mm.); fruits globose-elliptical about 
10 x 9 cm., glaucous when ripe. Leaves beneath with the quaternary 
nerves, minor veins, and areoles covered by minute tomentum of minute, 
thin, white, stellate hairs 2. T. sylveatre 
2. Filaments 3-antheriferous; inflorescences on the trunk, multiflorous; flowers 
rather large (sepals 10-12 x 3.6-4 mm.). 
3. Leaves with glabrous primary and secondary nerves beneath or sub- 
glabrous with very scattered mediocre stellate hairs and sparse callose 
spots; fruit globose-ellipsoid, 10 x 8 cm., without ribs, shortly tomentose, 
yellowish when ripe 3. T. speciosum 
3. Leaves softly velutinous beneath, the nerves and veins with abundant 
long, thin, patulous stellate hairs; fruits ellipsoid, densely velutinous, 
with 5 very prominent ribs 4. T. vel utinum 
1. Leaves with glabrous nerves and veins beneath, or subglabrous with very 
sparse, mediocre stellate hairs, only the areoles covered with compact 
tomentum of minute, white, stellate hairs. 
CUATRECASAS—CACAO AND ITS ALLIES 453 
4. Flowers large: petal-lamina suborbicular 5.5-7 x 5-6.5 mm.; petal-hood 
5-6 x 2.5-3 mm.; staminodes lanceolate-subulate 10-12 mm. long; fila- 
ments 2-3-antheriferous; sepals 12-13 x 3-4 mm.; petal-hood puberulous; 
fruit ellipsoid, obtusely pentagonal, attenuate at apex, umbilicate at base, 
10-11 x 5-5.5 cm. Leaves coriaceous, broadly ovate or ovate-oblong. 
5. T. glaucum 
4. Flowers smaller: petal-lamina suborbicular, orbicular or elliptic, 2.5-4 mm. 
long; staminodes 6-9 mm. long; filaments 2-antheriferous; sepals 8-10 x 
3 mm.; petal-hood hirtellous pubescent; fruit ellipsoid-oblong, obtusely 
pentagonal, abruptly narrowed at apex, umbilicate at base, constricted 
or not above the base, 12-25 x 5.2-8 cm 6. T. bernouillii 
KEY TO SECTION TELMATOCAKPUS 
1. Leaves regularly penninerved; inflorescences on trunk and branches; pedun- 
cles 5-25 mm. long; pedicels 7-8 mm. long; fruit peduncles 2-3 cm. long; 
sepals stellate-tomentose; petals pilose above, narrowed in the lower third, 
the apex blunt or emarginate; stami node-base with very short, thick 
hairs; ovary ovoid-ellipsoid, tomentose; fruit, when ripe, ovoid, attenuate 
at apex, slightly 5-costate, alveolate, the epicarp densely appressed tomen- 
tose, 7.5-11 x 7-9 cm  8. T. gileri 
1. Leaves at base 3-nerved, the two lateral basal nerves ascending at an acute 
angle, the other 2 or 3 pairs of secondary nerves remotely higher; inflores- 
cences axillary on young branchlets; peduncles 0.5-1 mm. long; pedicels 
0.5-1 mm. long; fruiting peduncles 4-8 mm. long and thick; sepals with 
sparse stellate hairs; petals glabrous, gradually attenuate to the base, the 
apex acuminate, the acumen acute, 2-dentate; staminode-base with rather 
thick, long, flexuose hairs; ovary pyriform, glabrous or sparsely granulate, 
rarely sparsely stellate-pilose; fruit, when ripe, ellipsoid-globose, conspicu- 
ously 10-costate, reticulate-alveolate, usually 6.5-7 x 6-6.5 cm. 
9. T. microcarpum 
KEY TO SECTION GLOSSOPETALUM 
1. Inflorescences born on the trunk and main branches. Flowers large; stami- 
nodes obovate-oblong or oblong-spathulate, 9-11 mm. long, erect in anthesis; 
leaves large (20-54 x 8-30 cm.), coriaceous, obtuse at both ends, strongly 
nerved and tomentose beneath; stipules coriaceous, persistent; calyx 5- 
lobate, cupular at base reflexed. 
2. Petal-lamina and staminodes yellow. 
3. Fruit ellipsoid-oblong or ovoid-oblong, attenuate at both ends, umbilicate, 
obtusely pentagonal, 25-35 x 10-12 cm.; leaves oblong-elliptic or ovate- 
elliptic, subvelvety beneath with minute, stellate, flexuous, white hairs 
crowded on the areoles and minor veins, and other equal or slightly 
larger hairs on the other nerves; stipules oblong-lanceolate, subacute; 
petal-lamina subdeltoid-spatulate; staminodes oblong-obovate. 
UT nirmnlinae 
3. Fruit ovoid-ellipsoid or ellipsoid, smooth, rounded at both ends or slightly 
attenuate at apex, 18-22 x 9-11 cm.; leaves oblong-elliptic or ovate- 
elliptic, tomentose beneath with minute, stellate, flexuous, white hairs 
crowded on the areoles and minor veins, and other larger, thicker, fer- 
ruginous hairs copious on the other nerves; stipules ovate or ovate- 
oblong, obtuse; petal-lamina subtriangular-spatulate; staminodes 
oblong-obovate  12. T. stipulatum 
454 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
2. Petal-lamina and staminodes red. Fruits smooth (not ridged). 
4. Fruit ellipsoid-oblong, rounded at apex, umbilieate, 16-40 x 6-11 cm.; 
leaves obovate-oblong, densely covered beneath with minute, stellate, 
white, intricate hairs on the areoles and minor veins, other mediocre, 
stellate, ferruginous hairs sparse or copious on nerves, and larger ones 
with longer rays usually copious on the major nerves; stipules lanceo- 
late; petal-lamina subtrapezoid, attenuate at base; staminodes 
oblong-obovate  14. T. simiarum 
4. Fruit ellipsoid-ovoid, rounded at base, obtuse at apex, 19-20 x 10-11.5 
cm.; leaves oblong-elliptic or ovate, often rugose, densely covered 
beneath with minute, stellate, white, intricate hairs on areoles and 
minor veins, and other larger ferruginous hairs with longer, spreading 
rays abundant on nerves; stipules ovate, rather obtuse; petal-lamina 
obovate-deltoid, attenuate at base; staminodes obovate-oblong. 
13. T. chocoense 
1. Inflorescences small, axillary on foliose branches. 
5. Petal-lamina and staminodes yellow; staminodes oblong-obovate, erect. 
Calyx trilobate, reflexed; leaves thin, subcoriaceous, subobovate-oblong 
or elliptic-oblong (9-25 x 3-9 cm.), attenuate at both ends, acute, cinere- 
ous beneath with minute, stellate, whitish, intricate hairs and larger ones 
with longer, patulous rays on nerves; stipules membranaceous, subulate, 
deciduous; petal-lamina subobovate-spatulate, bilobate, emarginate; 
fruit ellipsoid-elongate, 5-angulate, more or less irregularly tuberculate, 
10-18 x 6—9 cm.   10. T. angustifolium 
5. Petal-lamina and staminodes red; staminodes curved, spreading in anthesis. 
6. Young branchlets, petioles, and buds covered with a woolly-floccose, 
ochraceous or tawny, deciduous tomentum. Adult leaves glaucous 
or glauco-cinereous, monotrichous beneath, with the principal and ter- 
tiary nerves glabrous, glossy, reddish punctate, the areoles and reticulum 
whitish tomentose by minute stellate hairs. 
7. Flowers large; calyx (14-15 mm. long) 3-lobate; staminodes very acute, 
lanceolate, spreading (9-13 mm, long); petal-lamina trapezoid- 
elliptic, thick, dark red; leaves firm-coriaceous; stipules subcoriaceous, 
persistent; fruits ellipsoid, 16-25 x 10-12 cm., rounded at both ends, 
smooth 15. T. grandiflorum 
7. Flowers smaller; calyx (about 7 mm. long) 5-parted; staminodes oblong- 
elliptic, rounded at apex (6 x 2.8-3 mm.); petal-lamina suborbicular, 
red, rather thick; leaves thin-ch&rtaceous, very asymmetrical at 
base; stipules membranaceous, linear-subulate, deciduous; fruit 
ellipsoid-obovoid, acute, granulate-tuberculate, 5-7 x 3-4 cm. 
16. T. obovatum 
6. Young branchlets, buds and petioles hirsute, or short-tomentose. Calyx 
5-parted. 
8. Young branchlets, buds, and petioles densely hirsute or hirsute- 
tomentose; leaves beneath tomentose-hirsute (especially in young 
plants), ferruginous or ochraceous by spreading long-radiate hairs 
on the nerves and veins, and with minute, white, intricate ones 
covering the surface. 
9. Pedicels 5-10 mm. long; peduncles 5-10 mm. long; inflorescences 
loose; sepals about 10 mm. long; petal-lamina and staminodes not 
ciliate; fruit ellipsoid-pyriform, smooth .... 20. T. sixiuosum 
CUATRECASAS—CACAO AND ITS ALLIES 455 
9. Pedicels up to 1 mm. long; peduncles up to 6 mm. long; inflorescences 
compact, glomerate; sepals about 7 mm. long; petal-lamina and 
staminodes ciliate; fruit unknown . . . . 21. T. cannmanense 
8. Young branchlets, buds, and petioles densely subappressed-tomentose; 
leaves cinereous or ferruginous beneath with minute, stellate, whitish, 
intricate hairs covering the areoles and the smallest reticulum and 
mediocre, thicker, ferruginous hairs, copious or scattered on the 
nerves. 
10. Bracteoles narrowly linear. 
11. Staminodes scarlet, lanceolate, acute or subacute, 6-7.5 x 2 mm.; 
petal-lamina scarlet, orbicular or subrounded, thick, 2-2.5 s 
2.2-3 mm.; ovary glabrous and smooth or very sparsely granu- 
lar; fruit ellipsoid or oblong-ellipsoid, often slightly attenuate 
at base, 7.5-11.5 x 5-6.6 cm., the pericarp tomentose, smooth, 
3-4 mm, thick when dry  17. T. subincanum 
11. Staminodes brownish red, obovate-oblong, rounded or subspatu- 
late at apex, 5-5.5 x 2-3 mm.; petal-lamina obovate-subrhom- 
bic, 4-5 x 4 mm,; ovary densely tomentose; fruit ellipsoid, 
rounded both ends, 7x4 cm 18. T. hylaeum 
10. Bracteoles 3, orbicular, cochlear, embracing the single bud; stami- 
nodes brownish red, obovate-oblong, rounded at apex, 6-7.5 x 
3,5-4.5 mm.; petal-lamina oblong-elliptic or oblong-obovate, 
5-7 x 3-5 mm.; ovary densely tomentose; fruit ellipsoid, rounded 
at apex, constricted above the base, 8-10 x 4.5-6 cm., the peri- 
carp, when dry, 1-1.8 mm. thick, the epicarp 1 mm. thick, 
fragile at maturity  19. T. nemorale 
Artificial key to the species 
1. Leaves glabrous or puberulous beneath. 
2. Leaves firmly coriaceous, regularly penninerved, with 10-14 secondary 
nerves each side, usually 20-30 x 7-10 cm,; fruits glabrous, ovate-oblong, 
more or less pentagonal or decagonal, with carnose relatively thick peri- 
carp; stamens 2-antheriferous; staminodes linear-subulate, erect in bud, 
red; petal-lamina spatulate, stipitate, yellowish; jorquette with 5 
branches; growth below jorquette. Flowers on trunk and on branchlets. 
7. T. cacao 
2. Leaves chartaceous with 3-6 pairs of secondary nerves; fruits smaller, 
ovoid or subglobose, with hard-costate reticulate tomentose pericarp; 
stamens 3-antheriferous; staminodes subulate-subflagelliform, flexuous 
in bud; petals lacking laminae or ligular appendages; jorquette with 3 
branches; growth above jorquette. 
3. Leaves 5-20 x 1.5-8 cm., regularly penninerved with 5 or 6 nerves on 
each side, puberulous beneath, the midrib tomentulose; peduncle 
5-25 mm, long; pedicels 7-8 mm. long; ovary tomentose; fruit ovoid, 
slightly 5-costate and reticulate, 7.5-11 x 7-9 cm., fruiting peduncle 
2-3 cm.; flowers on trunk and on branchlets. Sepals tomentulose. 
ST irilArS 
• JL * giicri 
3. Leaves 6-16 x 2-7 cm., 3-nerved at base, the two lateral-basal nerves 
ascending, quite distant from the other 2 or 3 pairs of secondary 
nerves, glabrous or minutely and sparsely puberulous beneath; pe- 
duncle 0.5-1 mm. long; pedicel 0.5-1 mm. long; ovary glabrous or 
sparsely granulate; fruit ellipsoid or subglobose ellipsoid, strongly 
680-695—64 6 
456 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
10-costate and reticulate 6.5-7 (-9) x 6-6.5 cm.; fruiting peduncle 
4-8 mm. long; flowers only on branchlets ... 9. T. microearpum 
1. Leaves densely stellate-tomentose beneath. Jorquette 3-branehed. 
4. Leaves thin, chartaceous, tomentose-cinereous beneath. Flowers on ter- 
minal, leafy branchlets. 
5. Leaves ovate or ovate-oblong, subcordate or cordate at base, cinereous 
or ochraceo-cinereous (more or less silvery) beneath, the hairs uniform, 
minute, stellate, covering the surface and nerves; stamens 2-antherif- 
erous; petal-lamina subsessile; growth below jorquette, Dichotomous 
inflorescences on branches; fruit large, ellipsoid with thick-woody, 
strongly costate-reticulate and lacunose pericarp; flowers small, red 
dish; staminodes thick-linear, obtuse 1. T. bicolor 
5. Leaves oblong, elliptic-oblong, lanceolate-oblong, regularly pinnate- 
nerved, obtuse or cuneate at base, tomentose-cinereous beneath; 
stamens 3-antheriferous; petal-lamina stipitate; growth above 
jorquette. 
6. Leaves obovate-elliptic or obovate-oblong, obtuse and very asym- 
metrical at base with 5-7 nerves each side, 7-35 x 3-13 cm.; homo- 
trie hous, covered by minute, dense, white, stellate hairs beneath, 
except for the glabrous red-punctate main veins; young vegetative 
parts with a floccose, lanate, ochraceous, deciduous indument; fruit 
small, ellipsoid-obovoid, tuberculate-warty, 5-7 x 3-4 cm.; calyx 
5-1 obate. Flowers small; staminodes petaloid, red. 
16. T. obovatum 
6. Leaves elliptic-lanceolate or oblanceolate, rather oblong, slightly 
asymmetrical at base, heterotrichous beneath with dense, white, 
minute, stellate hairs and longer, patulous or subpatulous, pale 
ochraceous ones on the main nerves; young vegetative parts mi- 
nutely tomentose; fruit large; calyx 2- or 3-lobate. 
7. Leaves subobovate-oblong or elliptic-oblong or oblanceolate, acute, 
9-25 x 3-9 cm., with 6-8 secondary nerves on each side; petal- 
lamina and erect staminodes yellow; fruit ellipsoid-oblong, ir- 
regularly sulcate tuberculate, 10-18 x 6-9 cm. 
10. T. angustifolium 
7. Leaves elliptic-oblong or sublanceolate, 10-25 x 3.5-8.5 cm., with 
9-12 secondary nerves each side; petal-lamina small, narrow, 
red; staminodes red-purplish, very broad, reflexed, covering the 
stamens and petals; fruit ellipsoid-oblong, constricted above the 
base and at tho top below the apex, 10-20 x 6-8 cm. 
22. T. mammosum 
4. Leaves coriaceous, firmer and more markedly nervose-reticulate than in 
the species above. 
8. Leaves ovate-oblong or elliptic-oblong, long-caudate, with curved, as- 
cending secondary nerves, softly velvety or apparently glabrous and 
shining beneath; growth below the jorquette; petal-lamina sessile; 
staminodes lanceolate or subulate, erect in bud. 
9. Leaves more or less bullate, softly velvety beneath, heterotrichous 
with a layer of dense, minute, white, stellate hairs and longer, 
thin-rayed, patulous, stellate hairs on the nerves. Flowers purplish 
red on trunk; fruit ellipsoid with 5 protuberant ribs, 8-9 x 6-6.3 cm. 
4. T. velutinum 
CUATRECASAS—CACAO AND ITS ALLIES 457 
9. Leaves flat, with practically homotrichous, cinereous or whitish 
1 
indument beneath, of minute, white, stellate hairs; larger hairs 
very rare, the major nerves glabrous. 
10. Leaves beneath with glabrous major nerves, the quaternary and 
minor veins Stella te-tomentulose, the areoles densely whitish 
tomentose. 
11. Inflorescences small, on leafy branchlets; flowers small, brownish 
red; stamens 2-antheriferous; fruit ellipsoid-globose, glaucous, 
10 x 9 cm  2. T. sylveatre 
11. Inflorescences large, on the trunk; flowers purplish red, larger than 
above; stamens 3-antheriferous fruit globose-ellipsoid, 10 x 8 
cm., yellowish 3. T. speciosum 
10. Leaves beneath with completely glabrous veins, only the smallest 
veins of the reticulum sub glabrous with scattered mediocre 
hairs, the areoles with very appressed tomentum of minute, 
white, stellate hairs, the leaf surface with a glabrous appearance. 
12. Flowers 12-13 mm. long; petal-lamina suborbicular 5.5-7 x 
5-6.5 mm. long; staminodes Lanceolate-subulate, 10-12 mm. 
long; stamens usually 3-antheriferous, also 2-antheriferous; 
fruit ellipsoid, obtusely pentagonal, 10-11 x 5-5.5 cm. 
5. T. glaucum 
12. Flowers 8-10 mm. long; petal-lamina elliptic, suborbicular or 
orbicular, 2.5-4 mm. long; staminodes 6-9.5 mm. long; sta- 
mens 2-antheriferous; fruit ellipsoid-oblong, more or less 
pentagonal; 12-25 x 5.2-8 cm 6. T. bernouillii 
8. Leaves broad or oblong with regularly spreading, pinnate, secondary 
nerves, cinereous or ferruginous tomentose beneath, with prominent 
venation; growth above the jorquette; petal-lamina pedicellate or 
very reduced; staminodes broadly oblong. Stamens 3-antheriferous. 
13. Inflorescences on leafy branches. 
14. Leaves beneath monotrichous, glaucous, covered by minute, white, 
intricate, stellate hairs, the main nerves glabrous with scattered 
reddish, callous spots; young vegetative parts with ferruginous, 
floccose-lanate, deciduous indument; calyx trilobate; fruit large, 
ellipsoid, smooth, 16-25 x 10-12cm.; stipules persistent. Flowers 
dark red  15. T. grandiflorum 
14. Leaves beneath heterotrichous, with minute, white, densely intri- 
cate, stellate hairs covering the surface, and larger, thicker, 
reddish or ochraceous stellate hairs on the veins; calyx 5-lobate; 
fruits ellipsoid, smooth, 7-11.5 x 4-6.6 cm.; stipules caducous. 
15. Young branchlets hirsute or hirsute-tomentose; leaves with long, 
spreading radiate hairs beneath. 
16. Pedicels 5-10 mm. long; peduncles 5-10 mm. long; sepals 
about 10 mm. long; staminodes and petals not ciliate. 
20. T. sinuosum 
16. Pedicels almost lacking (up to 1 mm. long), peduncles up to 
6 mm. long; sepals about 7 mm. long; staminodes and petals 
ciliate 21. T. canumanense 
15. Young branchlets and leaves beneath subappressed-tomentose. 
17. Bracteoles broadly ovate or orbicular. Staminodes obovate- 
oblong, rounded at apex; petal-lamina oblong, obovate spat- 
ulate; ovary densely tomentose 19. T. nemorafa 
17. Bracteoles linear. 
458 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
18. Staminodes scarlet, lanceolate, acute, or subacute; petal- 
lamina scarlet, orbicular or suborbicular; ovary glabrous 
or very sparsely granular 17. T. subincanum 
18. Staminodes brownish red, obovate-oblong, rounded or sub- 
spatulate at apex; petal-lamina obovate, subrhombio; 
ovary densely tomentose 18. T. hylaeum 
13. Inflorescences on trunk and branches. Leaves usually large and thick- 
coriaceous, strongly nervose beneath; stipules persistent. 
19. Flowers yellow. 
20. Indument of leaves beneath sub velvety, the minute stellate hairs 
on veins slightly larger than those of the surface; stipules oblong- 
lanceolate, subacute; fruits ellipsoid-oblong, obtusely pentag- 
onal narrowed at apex, 25-35 x 10-12 cm. . 11. T. cirmolinae 
20. Indument of the leaves beneath, tomentose, the hairs on the 
veins ferruginous, larger than those of the surface; stipules 
ovate, obtuse; fruits ovoid-ellipsoid, rounded at both ends, 
smooth, 18-22 x 9-11 cm 12. T. stipulatum 
19. Flowers purple-red. 
21. Leaves obovate-oblong with three kinds of hairs beneath (minute, 
mediocre, and longer); stipules lanceolate; fruit ellipsoid- 
oblong, rounded at apex, 16-40 x 6-11 cm. . . 14. T. simiarum 
21. Leaves oblong-elliptic or ovate, usually rugose, with two kinds of 
hairs beneath (minute and larger); stipules ovate, rather 
obtuse; fruit ellipsoid-ovoid, rounded at base, obtuse at apex, 
19-20 x 10-11.5 cm. .  13. T. chocoense 
Section I* Rhytidocarpus 
Theobroma sect. Rhytidocarpus Bernoulli, Uebers. Art. Theobroma 9. 1869. 
Sect. Eutheobroma subsect. Rhytidocarpus (Bernoulli) Pittier, Rev. Bot. Appl. 
10(110): 779. 1930, 
Petal-lamina very shortly stipitate, subsessile. Petal-hood 1-nerved. 
Staminodes linear-oblong, obtuse, thick, erect in aestivation. Fila- 
ments 2-antheriferous. Fruits subglobose-ellipsoid with hard peri- 
carp, strongly costate and reticulate-nerved, minutely tomentose, 
the mesoearp thick-woody, very hard. Cotyledons epigeous at ger- 
mination. Leaves beneath, appressed stellate-tomentose. Primary 
1 eaves palmatinerved, regular leaves subpalmatinerved, the base 5-7 
Served. Inflorescences axillary or extra-axillary on leafy branches. 
Sympodial growth of stem by orthotropic, adventitious, lateral- 
Sub terminal shoots. Primary branches tern ate, deciduous in age, 
leaving a naked stem; leafy crown lax, flat. Secondary branching 
dichotomous. 
Type species.—Theobroma bicolor Humb. & Bonpl. 
A single species known. 
1. Theobroma bicolor Humb. & Bonpl. Figures 2, 5, 9, 12, 18, 35; Map 3 
Theobroma bicolor Ilumb. et Bonpl. PL Aequin. 1:104, pL SO. 1806; H, B. K. 
(1823) 317; Triana & Planch. (1862) 208; Bernoulli (1869) 9, pi. 4; Schu- 
mann in Mart. (1886) 73; Jumelle (1899) 21, figs. 10, 11; Preuss (1901) 
CUATRECASAS—CACAO AND ITS ALLIES 459 
Figure 12.—Leaves of Tkeobroma, X H* Ai bicolor, from orthotropic branches (Dawe 83), 
from above; b, base of same from underside; c, tricolor from plagiotropic (current) 
branches (Kill. & Smith 30006); D, speciosum van coriaceum (Huber 1567); e, velutinum 
(Beaoist 516). 
460 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
251, 255, pis. 5, 4; De Wildeman (1902) 94; Huber (1906a) 274; Standley 
(1923) 808; Ducke (1925) 132; (1940) 269, pi. £; (1954) 13; Standley 
(1937) 687; Chevalier (1946) 276; Standl. & Stey. (1949) 422; Holdridge 
(1950a) 3; Addison & Travarea (1951) pis. 10, IS, jig. 8; Baker, Cope & al, 
(1954) Jig. 9; Cuatrecasas (1956) 652; Le6n (1960) 313, 315 jig. 
Theobroma ovatifolia Moc. & Sess£ ex DC. Prodr. 1:485. 1824; Icon. PI. 
Mex. DC. pi. US, 
Cacao bicolor (Hurab. & Bonpl.) Poir. in Lam. Encycl. M6th. Suppl. 2:7. 
1811. 
THbrotna bicolor (Humb. & Bonpl.) Cook, Journ. Washington A cad. Sci. 
5:288. 1915; Contr. U.S. Nat. Herb. 17(8), pis. 46, 47, 48, 49, SO, 52, 
64, 1916. 
Theobroma cordata Rufz & Pav6n, Fl. Peruv. Chil. vol. 6, ined. 
Map 3.—Geographical distribution of Tkeobroma bicolor, based on specimens mostly from 
planted trees. 
Types.—Humboldt <& Bonpland, Colombia (P). MociHo <& Sess6, 
Mexico (of T. ovatifolia). 
Commonly a small tree 3-8 m. tall, attaining in high forest a height 
of 25-30 m,, with rather narrow crown; sympodial growth by lateral, 
subterminal, upright shoots; trunk erect with light bark and white 
wood; primary branches ternate, dichotomous, spreading; young 
branchlets often horizontal or pendulous, more or less flexuous, 
subterete, densely and appressed cinereous tomentose with minute 
CUATRECASAS—CACAO AND ITS ALLIES 461 
stellate hairs; older branches glabrate, smooth, gray; stipules oblong- 
lanceolate, 5-8 mm. long, 1.2-2 mm. broad, minutely appressed- 
tomentose, more or less persistent. 
Leaves subpalmatinerved, firmly chartaceous, green above and 
silvery greenish or silvery cinereous, sometimes pale ochraceous 
beneath; petiole rather thick, subterete, rigid, minutely appressed- 
toinentose, 12-25 mm. long, transversely rimose when old; blade 
oblong-ovate or elliptic-ovate, more or less deeply cordate or emargi- 
nate, asymmetrical at base, attenuate, abruptly acuminate at apex, 
entire or rarely sinuate at the upper margin, 12-34 cm, long, 6-18 
cm. broad, the acumen triangular, 6-12 mm. long, glabrous above or 
with scattered stellate or furcate hairs, green or when dry, pale 
brownish, the main nerves noticeable, the lesser slightly conspicuous, 
cinereous beneath, covered with a dense layer of intricate, white, 
sericeous, stellate hairs, at base 5-7-nerved, the thicker costa and 2 or 
3 main nerves on each side strongly prominent, the interior, basal nerves 
upright, ascending, the 1 or 2 exterior basal pairs arched, spreading, 
thinner, on the % upper part with about 4 secondary nerves each side, 
prominent, ascending, near the margin curved and vanishing, the 
tertiary transverse nerves prominent, the lesser prominulous veins 
minutely reticulate; leaves of young upright (orthotropic) shoots 
larger, symmetrical, long-petiolate, the blades broadly ovate, deeply 
cordate, more markedly palmatinerved, 30-50 cm. long, 21-36 cm. 
broad, the petiole 10-38 cm. long, thickened-pulvinate at both ends. 
Inflorescences axillary or extra-axillary on leafy, j uvenile branchlets, 
usually 3-6 cm. broad, with very short axis and divaricate, dichasial 
and cincinnate branching; branchlets and pedicels angulate, densely 
ochraceous or cinereous tomentose; peduncles very short, supporting 
an articulate pedicel subtended by a bracteole; pedicels erect, to- 
mentose, 3-6 mm. long; bracteoles lanceolate, sub acute, rather thick, 
more or less curved, densely and minutely tomentose, 1.5-2.5 mm. 
long, 0.6-1 mm. broad; buds oblong-ovate, subacute, slightly 5- 
angulate, densely adpressed and minutely cinereous or ochraceous 
tomentose. 
Sepals lanceolate or ovate-lanceolate, acute, shortly connate at 
base, spreading and more or less curved-indexed, 5-6 mm. long, 2-2.5 
mm. broad, 3-nerved and sparsely pilose inside, glandular at base, 
subappressed stellate-tomentose and reddish outside, the margin 
minutely whitish tomentellous. 
Petal-hoods 2-2.5 mm. long, 1-1.2 mm. broad,submembranaceous, 
whitish rosy, or reddish with darker midrib, oblong-obovate-elliptic, 
rounded-cucullate, emarginate, auriculate and incurved apex, glabrous 
with a thick trifurcate midrib inside, hirtellous-pubescent with a de- 
pressed tomentellous midrib outside; lamina rather carnose, red, 
462 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
brownish red or purplish, ovate, rounded at apex, abruptly contracted 
at base in short nail, articulate to the claw, hirtellous pubescent, 1-1.2 
mm. long, 0.8-1 mm. broad, the nail 0.2-0.3 mm. long, erect in bud. 
Androecium tube 1.5-1.8 mm. long; staminodes 5 mm. long, brown- 
ish red, usually lighter red toward the base with whitish margin, 
carnose, linear-oblong, obtuse or subacute, slightly narrowed at base, 
copiously covered with minute, thickish, patulous hairs, erect in bud, 
3.5-4.5 mm. long, 0.6-0.8 mm. wide; filaments compressed, curved, 
reflexed, glabrous, about 1-1.5 mm. long, shortly 2-furcate, 2-anther- 
iferous; anther lobes ellipsoid, 0.2-0.3 mm. long; ovary obovate-oblong, 
sharply 5-costate, greenish white, velutinous-tomentose, 1.8-2 mm, 
long, 1-4 mm. broad; styles whitish, about 1.7 mm. long, united, rigid. 
Fruits subglobose-ellipsoid, oblong-ellipsoid, or ovoid-ellipsoid, 
15-20 (10-25) cm. long, 9-12 (-15) cm. broad, green, when ripe yel- 
low or brownish; pericarp hard, strongly 10-costate, the commissural 
ribs thick and elevated, the 5 alternate similar but diminishing in 
thickness towards the apex, the deep furrows reticulate (ligno-nerved), 
deeply lacunose; pericarp composed of three layers: 1) carnose endo- 
carp about 2 mm. thick, 2) woody, very hard, ribbed and nerved 
mesocarp, 3-6 mm. thick, 3) firm, carnose becoming coriaceous epi- 
carp 1-2 mm. thick with an outer, densely stellate-pilose epiderm, 
the whole pericarp about 7 mm. thick on the furrows and 14-15 mm. 
thick on the ribs; pulp surrounding seeds fibrose, yellowish, sweet, 
scenty; seeds arranged in 5 rows, complanate, ovoid-amygdaliform, 
16-30 mm. long, 14-23 mm. wide, 8-13 mm. thick; embryo white; 
germination epigeous. 
Theobroma bicolor is unique in the genus, easy to recognize by its 
small, lax crown of few whorls of horizontal, dichotomous branches 
topping a naked stem, and by its leaves, fruits and flowers. The 
large papery, firm leaves are whitish silvery beneath and those of 
the upright shoots are larger, palmatinerved, cordate, and very long 
petiolate. The flowers are small and pale red generally. The large 
fruits are strongly ribbed and nerved, with a hard, woody, carved 
shell; they keep the green color until ripening, when they become 
yellow or brownish, falling from tree. 
Common names.—In English: Patashte. Mexico: Patashte, 
patashtle, pataste, petaste, patatle, petaxte, patasht, pataste de sapo, 
pataste simarron, cacao malacayo, cacao bianco. Guatemala: Patashte, 
pataxte, patasht, balamati, balam (KekcH), pec (PoJconchi). Costa 
Rica: Pataste, pataiste, skar-ub (Bribri), uerba (Terraba), scarbo 
(Bribri), carvu (Kahekara), sapar<5n (Estretta), erefa (Quatuso). 
PanamA: Pataste, cu-lu-hu (CJioko). Colombia: Bacao (general), ca- 
CUATRECASAS—CACAO AND ITS ALLIES 463 
cao silvestre, cacao marraco, and marraco (in Caquet&). Ecuador: 
Patas, cacao bianco. Peru: Macambo, majambo, najambu. Brazil: 
Cacau do Peru (Belem), cupuassti, cupua-I, cacau baftj, cacao bravo. 
The Anglo-Colombian Cocoa Expedition (Baker, 1953) recorded 
the following names: he£-a (Maku), (Piraparan&, Taraira); a5 (Ma- 
kuna) (lower Piraparan&, Popeyaca); la-na-pee-ta-ma-ca-la-chu-na-ni 
(Yakuna) (Miritiparan&); ha-ha (Tanimuka) (GuacayS.); maraca 
bacao (Choc6). 
Uses.—The pulp is frequently eaten or used by natives to prepare 
refreshments although its flavor is not very attractive. The seeds 
are used in many places like those of cacao, giving a chocolate of 
inferior quality; it is also locally used to manufacture pastry and 
candy. The seeds of T. bicolor have been commercially mixed some- 
times with those of cacao. They are poor in theobromine but they 
have a great proportion of a good quality cocoa butter. In Guate- 
mala the seeds are known in commerce as "tiger" 'fwariba" or "pa- 
tashte" cacao (Standley). 
According to Llano (1947), in Colombia this species has been tried 
for grafting but the bark heals with difficulty. 
The hard shells of the pod are used locally, as containers like those 
of mate or tviuma (Crescentia cujete). According to Tafalla it is 
called "cacao de Castilla" in the Ecuadorian region of Uchiza, where 
the shells of the pods are smoothed to be used as bowls. 
Distribution.—Theobroma bicolor is widespread in cultivation 
throughout all humid tropical America, from southern Mexico to 
Bolivia and Brasil. It is never cultivated extensively, but a few trees 
can always be seen where cacao is cultivated, and usually also in 
backyards of tropical farms and in secondary growth. It is fre- 
quently found also subspontaneous in more or less open thickets. I 
have always seen the species in cultivation only. Its true native place is 
uncertain. Probably it originated in Central America where it is 
said to be found in primary forests. Miranda (I.e.) writes "se en- 
cuentra en acaguales de las selvas alt as" in Chiapas; Steyermark 
found it in dense forests of the region of Ixcan (Huehuetenango) in 
Guatemala. Another possible original region is the eastern region of 
Peru and Ecuador. Ynes Mexia recorded it from dense forests in 
the Napo-Pastaza river basin. Llewellyn Williams found it spon- 
taneous in secondary growths in Lore to. But thus far I have no 
assurance of a place where it is incontestably native. 
MEXICO: Herbarium Sessfi et Mocifio No. 3620, preserved at the Madrid 
Botanical Garden "Plantae Novae Hispaniae a Sess6, Mocifio, Castillo et Mal- 
donado lectae (1787-1795-1804)-Theobroma Patastle Ie. N." (MA, lectotype of 
T. ovcUifolium; BM, F, G, isotypes); the MA specimen has foliage, inflorescences 
and flowers and may be considered the holotype of Theobroma ovatifolium Mocifio 
464 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
et Sess6 ex DC.; I select it as the lectotype; it is photograph FM No. 48412. 
The Chicago specimen is a duplicate of this and has only one leaf; the very good 
British Museum specimen, and two at Geneva are certainly isotypes in spite of 
not bearing the name of Scss6 and Mocifio and having "Theobroma Palastle, 
N.E." as only information; some specimens have the indication "Herb. Pav6n," 
having been sent by Pav6n from Madrid. The collections with the number 
3621 Sess€ et Mocifio et al. (MA) are paratypes and may be duplicates of 3690, 
having been numbered recently (in 1935); they include normal foliage and inflo- 
rescences and juvenile long-petiolate, broadly cordate leaves (FM photographs 
48414 and 48413). The photograph FM 30526 is from plate 485 of the Mocifio 
and Sess6 Flora Mexicana, No. 113 in De Candolle's copies. 
Oaxaca: Po chut la, Capital Rancho Vie jo; tree 15 m., flowers purple, "cacao 
malacayo," Feb. 1941, Reko C068 (F). 
Chiapas: Acagoyagua, Escuintla, "pataste de sapo," 22 VII 1947, Matuda 
16733 (F). Ibidem, "pataste simarr6n," fruit only, 6-7 x 4 cm., 28 VIII 1947, 
Matuda 16840 (F, MEXU). Esperanza, Escuintla, "pataste," cultivated, 11 
VIII 1947, Matuda 16690 (BR, F, MICH, MEXU, NY). Palenque, 100-150 
m. alt.; tree up to 25 ft. tall; commonly cultivated around Palenque for its fruits; 
fruits yellow; seed ground and mixed with cornmeal for making "pazoli"; flower- 
ing and fruiting in May; said to be common in the forests on the slopes above 
Chacamax River, V 1937, LI. Williams 9345 (F, Y). 
GUATEMALA: Without locality, "cacao," 10 V 1914, Davidson s.n. (US). 
Alta Vjsrapaz: Trece Aguas, cacao plantation at Cacao, "petaste" or "patashte," 
9 III 1914, Cook & Doyle 50 (US); Trece Aguas, near Finca Sepacuit^, "petaxte" 
IV 1905 (leaf from upright stem of young tree), Cook 4 (US). Ibidem, Chirujija 
Oxee, near Finca Sepacuit<5, "balamati," 25 V 1902, Cook & Griggs 756 (US). 
Cham a, 900 ft. alt.; tree 25 ft., flowers reddish, fruit large, hard shelled, "patashte" 
cultivated by Indians; used similarly to cacao, rather sweeter in flavor, 15 VI 
1920, H, Johnson 237 (F, US). Cubilquita, 350 m, alt., VI1901, von Tuerckheim 
7824 (GH, K, NY, US). Vicinity of Sibicte, 370 m. alt., small tree 25 ft. tall, 
leaves firmly membranaceous, deep green above, gray silvery beneath; flowers 
with grapy-purple calyx, "balam" (Kekchi), "patasht" (Spanish), cultivated, 
12 III 1942, Steyermark 44941 (F). 
Huehuetenango: Sierra de los Cuchumatanes, 150-200 m., between Ixcan 
and Rfo Ixcan; tree 30 feet tall, leaves membranaceous, papyraceous, dark green 
above, silvery green beneath; dense rich forest, 23 VII 1942, Steyermark 49317 
(F, US). Huehuetenango, 1800 m. alt., seeds in market brought here from the 
Pacific Coast, Standby 82446 (F). 
Santa Rosa: Region of Platanares, between Taxisco and Guazacap&n, 220 
m. alt,; wet forested quebrada, simple shrub 3-4 m, tall; escaped here, 3 XII 1940, 
Standley 79069 (F). 
Suchitepequez: Mazatenango, cultivated, III 1865, Bernoulli 94 (F, G, NY). 
Ibidem, III 1865, Bernoulli & Cario 3145 (GOET). 
BRITISH HONDURAS: Conservation Forests H. 2192/29 (F, GH, UC). 
Stann Creek Valley, Big Eddy Valley; tree 13 inch diameter, fruit 8 inch long, 
"pataste," "mountain cacao," 16 XII 1840, Gentle 3464 (F, GH, MICH, MO, NY, 
U). 
EL SALVADOR: Sonsonate, cultivated, "patashte," "pataste," 20 IV 22, 
Calderdn 23610 (F, GH, MO, NY, US); ibidem, "pataste," 1922, CalderAn 627 
(GH, NY, US). 
COSTA RICA: Peninsula Osa ad Golfo Dulcc, circa Puerto Gim&iez, ad litus; 
arbor 6-8 m., floribus purpurois, 15 IV 1930, Cudofontis 92 (WU). Vicinity of 
Guapilds (prov. Lim6n), 300-550 m. alt.; planted tree 25 ft., flowers dull red 12, 
CUATRECASAS—CACAO AND ITS ALLIES 465 
13 III 1924, Standley 37374 (US). Puerto Gim&iez de Osa and vicinity, 14IV1930, 
Brenes 12333 (Herb. Nac. C.R. 212) (F, NY). Edge of the road to Tuis, 650 m.; 
tree with spreading branches, "pataste" or "pataiste," XI1897, Tonduz 11304 (F), 
Tucurrique, grassland at Las Vueltas; tree 40 cm. diam., 15-20 m. high, 635 m. 
alt., "pataste," "pataiste," III 1899 Tonduz 13110 (G, P, US). Hacienda Balti- 
more (Lim6n), 10 m. alt., in a small plantation of this species, 8 VII 1949, Hold- 
ridge s.n. (TURRI), La Lola, planted, 6 XI 1961, Cuatrecasas & Paredes 26534 
(US). 
PANAMA: Bocas del Toro, in Laguna de Chiriquf and its neighborhood, 
Pope's Island, XI, XII 1885, Hart 158 (US). Dari6n, Headwaters ofRfoChica, 
500-750 ft.; tree 35 ft.; flowers dark red; cultivated by Choco Indians, "cu-lu-hu," 
Allen 4593 (G, MO, NY). Canal Zone, along Cafio Quebrado, 2 XII1914, PiUier 
6883 (GH, NY, US). 
TRINIDAD: L'Eranche Est, Sangre Grandre, 10 VII 1929, Boehlmer 12229 
(TRIN). Blue Basin, 2)4-3 miles distant, 21 IX 1928, Lange 12056 (TRIN). 
Grounds of I.C.T.A., River Estate Diego Martinez, field 19; fruits 19.3 x 13.3 cm., 
yellowish at maturity, specimen from seeds brought from Jinogoj6, Apaporis, 
Colombia, 31 VIII 1961, Cuatrecasas, Cope, & Bartley 25784T (US); same field; 
calyx pale red, hoods with darker midrib, petal-lamina very small, brownish red, 
staminodes brownish red, lighter reddish toward the base, with whitish margin, 
styles whitish, ovary greenish white, 1IX 1961, Cuatrecasas & Cope 25795 (US). 
Tree from seeds from La Pedrera, Colombia; fruits 16.2 x 14 cm., 31 VIII 61, 
Cuatrecasas, Cope, & Bartley 25787T (US). Field 2; fruits 15 x 10, 16 x 10.8, 
16.5 x 10, 18.2 x 11.2, peduncle 0.7-0.9 cm., 31 VIII 61, Cuatrecasas, Cope, 
& Bartley 25786T (US). 
SURINAM: Paramaribo Gardens, cultivated, VI 1910, Stockdale s.n. (K, U). 
COLOMBIA: Antioquia: Savaletas, 100-500 m., XII, Lehmann 7909 (K). 
To LIMA: Ibagu6, II1916, "cacao silvestre," Dawe 83 (US). 
Huila: Valle del Magdalena, Garzdn, IV 1845, Goudot s.n. (G, P). 
El Valle: Palmira, Granja Agrfcola, 900 m. alt. "bacao," Dugue Jaramillo 
1205 (F). Ibidem, X 1943, Llano s.n. (COL, F). Palmira, 925 m., cultivated in 
grounds of experiment station (said to have been brought from the Choc6); tall, 
erect tree, leaves hanging vertically, almost white beneath, flowers dark red with 
maroon staminodes, fruit oblong, green with the furrows strongly rugose, 29 X1944, 
Fosberg 21310 (NY, UC, US). Palmira; tree 4 m., "bacao," I 1947, Dugue Jara- 
millo 4403A (COL). Ibidem; tree 3 m., flowers red, "bacao," 3 XII 1947, Car- 
deflosa, Murgueitio, & Barkley 17C934 (COL, F). Cartago, Goudot s.n. (P, WU). 
Pacific Coast, Rio Calima, La Trojita, left side of river, 5-20 m. alt.; tree3 m., 
"bacao," 27 II 1944, Cuatrecasas 16526A (F, VALLE) (fruit only). Buenaven- 
tura (and Tumaco) 0-400 m., fl. perpetual, Lehmann 9021 (K, NY). 
Choc<S: "Nvelle. Grenade, Choc6," "bacao," Humboldt and Bonpland, s.n. 
(P, holotype). Bonpland, probably isotype, from Paris (F). Near Istmina, road 
to C6rtegui, in forest, 75 m. alt.; tree 6 m., flowers red, "bacao," 3 VII1944, Garcia 
Barriga 11178 (COL, US). Choc6 (i Barbacoas), 25 m. alt., "bacao," IV 1853, 
Triana 5333 (-3) (BM, COL). Ciudad Mutis, 27 X 1946, Romero Castafieda 
s.n. (COL). 
Nari&o: Pacific Coast, Amarales, "bacao," 1866, Triana s.n. (BM, BR, COL, 
G, K, NY, WU). 
CaquetA: Solano, 3 km. SE of Tres Esquinas, on Rio CaquetK, below mouth 
of Rio Orteguaza, wet tropical forest of Amazon basin; tree 8 m. high, 10 cm. 
DBH; dark brown, smooth, lichen patches, flowers red; watermelonlike fruit 18 
cm. long, 10-11 cm. broad, yellow, edible; Indians plant it; secondary lowland 
forest, river bank; possibly an escape from cultivation; several cultivated trees 
466 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
seen March 13 walk near Rio Caquetfi, 2 km. s. of Solano; "cacao marraco," 
6 III 1945, Little & Little 9598 (NY, US). 
Putumayo: Vicinity of Mocoa; small tree 4 m., sterile; apparently cultivated, 
17 III 1953, HoLliday & Cope T 77 (COL, TRIN, US). 
Vaupes: Rio Negro, near San Felipe, Cafto Mayabo, river level; tree 4-5 
years old, introduced on a house site, 27 X 1952, Baker 34 (COL, F, TRIN). 
Rfo Inirida, right bank below mouth of Cano Caribe; tree 25 m. high, 1 m. diameter 
at ground level, growing on slope just above high watermark; overtopping the 
surrounding forest; not thought to be wild. 23 I 1953, Barlley & Holliday T66 
(COL, TRIN, US). Rfo Vaupds, opposite confluence with Rfo Papurf, Yavarat£, 
Silesian Mission Sao Miguel; trees 3-4 years old from seed brought by Indian 
from the interior of Colombia, 20 II 1952, Bartley & Holliday T 47 (COL, TRIN, 
U, US). Cafio Umuna, Rfo Piraparand, river level; cultivated tree in Indian 
garden, 8 IV 1952, Baker & Cope 11a (TRIN). Jinogoj6, on Rfo Apaporis, small 
tree 15 ft. cultivated in Indian garden, 23 VIII 1952, Baker & Cope 2 (TRIN). 
Rfo Apaporis, Gino-Goj'6, between the rivers Piraparand and Popeyacri, 250 m. 
alt.; tree 4 m., leaves white below, 3-11 IX 1952 Garcia Barriga 14416 (US). 
Amazonas: Rfo Caquetd, La Pedrera, river level; tree 30 ft., presumed cul- 
tivated, 1 X 1952, Baker & Cope 2G (COL, F, TRIN, US). Rfo Igaraparand, 
vicinity of La Chorrera, 180 m. alt.; small tree, cultivated, "marraca," 4-10 VI 
1942, Schultes 3922 (COL, F). 
ECUADOR: Muller s.n. (K). Balao; arbor 40 m., in forest, flowers purplish, 
"cacao bianco," I 1892, Eggers 14244 (A, L, LE, M, US). Rfo Sucumbfos, be- 
tween Putumayo and Quebrada Teteyd, 260 m.t "cacao" (K of tin), 29 III 1942, 
Schultes 3471 (NY). 
Nafo-Pastaza: Between Tena and Napo; tree about 15 m. high, petals 
dark red, staminodia blackish brown, 5 I 1940, Asplund 10271 (S), NE of the 
province, Tiputini-Lagartococha, 20 1-5 II 1953, Fagerlind & Wibom 2371 (S). 
Cantdn Napo, near Tena, 400 m. alt., dense forest; tree up to 18 m. high, fruit 
called "Patas," size of small watermelon, the pulp being eaten and the seeds 
cooked or raw much appreciated, 2 II-VI 1935, Mexia 7214 (F, P, UC, US). 
BRAZIL: "Brazil, Dr. Martius," no data (G). Manaquiry forest; 15-30 ft., 
leaves white beneath, flowers dark purple. Spruce s.n. (K). 
Amazonas: Rio Negro, Barcellos, matta, cult.?, 27 VI 1905, Ducke 7202 
(BM, MG, US). Ibidem, Ducke 7202-B (BM, MG). "Prov. Rio Negro, in 
sylvis, Martius Obs. 28823, Iter-Brasiliensium 319, Martius [8C2, 863, 864, 865] 
(M). Rio Negro, Lagos, "cacao bravo," 5 VIII 1874, Traill s.n. (GH). Rio 
Negro, Schomburgk 870 p.p. (BM, G, GH, GL, K, OXF, P, US). Sa5 Antonio de 
Iga, capueira, "cupuassti," 3 VIII 1906, Ducke 7638 (MG). Municipio Sao Paulo 
de Olivcnga, basin of creek Belem; tree 40 ft. high, trunk 5 inch diam., planted by 
Indians 26 X-XII 1936, Krukoff 9019 (A, BM, F, G, IC, LE, MICH, MO, NY, P, 
US). Manaos, Campos Salles, 15 m., 20 VIII 1928, Luetzelburg 23895 (M). 
Manaos, Agricultural Experiment Station, 25 m. alt.; tree 30-35 ft., fruit oblong 
on ultimate branches (25 x 10 cm.), cult.?, 13 X 1929, KiUip & Smith 30006 (GII, 
NY, US), fruit collection 681 (US). Experimental Garden of Nord Brazil, culti- 
vated and wild in forest, "cacao dJAnta," 20 VIII 1928, Luetzelburg 23065 (M). 
Ega, "colitur circum Indorum villas, Oct. 1830," Pocppig 2746 p.p. (WU);"In 
Bras, tropica fl. Amazonas Oct. 1839," Pocppig 2746 pp. (GOET). Ega, Poeppig 
2746 p p. (F, LE). Teffti, forest, 29 VI 1906, Ducke 7397 (BM, G, MG, US). Fonte 
B&a, firm land, medium-sized tree, "cacau bafti," 28 III 1945, Fr6es 20625 (F, 
IAN, K, NY, US). Rio Jurud, Santa Clara, cultivated, "cupuagd" Baum, X 1900, 
Vie 5030 (BM, G, IIBG). Rio Jurud, Gaviao; flowers purple, "cupua-i," III 1875, 
Traill 60 (K, P). Rio Sapo, 21 II 1874, Traill 60 (K). "Ad oram meridion&lem 
CUATRECASAS—CACAO AND ITS ALLIES 467 
flum. Amazonum ad ostium flum. Solimoes," VI 1851, Spruce 1609 (BM, K, Mf 
P, WU). 
Ceara: Ceard, VIII-XI 1838, Gardner 870 (G). 
Fab A: Be 16m, Horto do Museu Goeldi, arvore 483; medium size tree, granate 
flowers, 10 X 1957, Cavalcanle 310 (MG, US). Belfim, cult., Pires <& Black 746 
(IAN). Bel6m, I.A.N. cult., "cacau do Perd, 6 XI 1952, Pires 4340 (IAN, NY). 
PERU: "Theobroma cordata del Peru, sp. nov.," Ruiz & PavSn (BM). "Peru- 
via, Herb-Pav<5n" (G). Perou, Pavdn 617 (G). 1909-1914 Weberbauer 6245. 
Loreto: Mishuyacu, near Iquitos, 100 m.; forest, tree 6 m. high, flowers 
dark violet and rose, clearing, "macambo," V-VI 1930, Klug 1523 (F, NY). 
Florida, Rfo Putumayo at mouth of Rfo Zubineta, 200 m.; forest clearing, tree 
4 m., flowers garnet, "macambo," III-IV 1931, Klug 2021 (A, F, GH, K, MICH, 
MO, NY, S, US). Parafso, upper river Itaya, 145 m.; "najambu," 1 X 1929, 
LI. WiUiams 3346 (F, US). Amazon River, Caballo-Cocha, 6 VIII 1929, LI. 
Williams 2149 (F, GH, US). Maynas, Poeppig (L), Poeppig 18 (BM). Marafi6n 
River from Iquitos to the mouth of Rfo Santiago at Pongo de Manseriche, ca. 
77°30' W., 1924, Tessmann 4079 (NY, S). 
Section 2. Oreanthes 
Theobroma sect. Oreanthes Bernoulli, Uebers. Art. Theobroma 7, 1869. 
Figure 4; Map 1 
Sect. Bubrotna subsect. Oreanthes (Bernoulli) Pit tier, Rev. Bot. Appl. 10(110) :779. 
1930. 
Petal-lamina sessile, large. Petal-hood 3-nerved. Staminodes 
linear-subulate or lanceolate, erect in estivation. Filaments 3- or 
2-antheriferous. Fruit more or less angulate or costate with coriaceous, 
tomentose pericarp, the endocarp rigid, thin-lignose. Cotyledons 
epigeous at germination. Leaves densely, minutely, stellate- 
tomentose beneath inside the reticulum. Inflorescences many- 
flowered, on the trunk or, in few cases, small and on leafy branches. 
Sympodial growth of stem by orthotropic, adventitious, lateral- 
sub terminal shoots. Primary branches ternate, deciduous when old, 
leaving an unusually long, naked stem. Leafy crown loose, flat. 
Secondary branching dichotomous. 
Type species: Theobroma speciosum Willd. ex Spreng. 
2. Theobroma sylvestre Mart. Figures 4, 9, 13, 16, 18; Map 4; Plate 1 
Theobroma sylvestre Mart, in Buchner, Repert. 35:24. 1830; Linnaea, Litt. 
Ber. 32. 1831; Bernoulli (1869) 14, pi. 7, jig. 1; Schumann in Mart. (1886) 
78 (as T. silvestre); Jumelle (1899) 34; De Wilde man (1902) 98. 
Theobroma Spruceana Bernoulli, Uebers. Art. Theobroma 9, pi. 8, fig. 1. 1869; 
Ducke (1925) 131; (1940) 270, pL 8; (1954) 13; Chevalier (1946) 276; 
Addison & Tavares (1951), pi. 6, fig. lt pi. 6, fig. A, pi. IB, fig. 4', Le6n (I960) 
318, 319, fig. 
Theobroma nitida Bernoulli, Uebers. Art. Theobroma 15, pi. 7, fig. 1869. 
Theobroma Martii Schum. in Mart. Fl. Bras. 12(3): 78. 1886; Jumelle 
(1899) 35; De Wildeman (1902) 98. 
Theobroma speciosum var. Spruceana (Bernoulli) Schum. in Mart. Fl. Bras. 
12(3):75. 1886; Jumelle (1899) 32, fig. 16; De Wildeman (1902) 95. 
468 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Figure 13.—Leaves of Theobroma, X a, speciosutn, from orthotropic and plagiotropic 
branches CUS—1031918); b, sylvestre (US—1693105); c, sylvestre (Martius, type). 
CUATRECASAS—CACAO AND ITS ALLIES 469 
Map 4.—Distribution of Theobroma sylvcsire, 0; T. bernouillii subsp. bernouillii 0; 
T. bernouillii subsp. asclepiadiflorum O; T. bernouillii subsp. capilliferum Q; T. 
velvtinum A. 
Types.—Martius, Brazil, Iter Brasil. 322 (in M, no. 891) (Photo 
F. M. 19644). Spruce 166, Brazil, Par&, Obidoa (of T. Spruceana). 
Martius, Brazil, Iter Brasil. 322 (in M, no. 890) (Photo F. M. 40709) 
(of T. nitida). 
Small-or medium-sized tree up to 12 m. high, sympodial growth by 
lateral, sub terminal upright shoots; primary branching ternate; 
terminal, leafy and floriferous branches dichotomous, terete, rugulose, 
dark brownish, the hornotinous ones pulverulent, stellate-tomen- 
tellous, soon glabrate or glabrous; stipules small, soon deciduous. 
Leaves distichous, firmly coriaceous; petiole short, robust, appressed 
stellate-tomentose, 5-10 mm. long; blades elliptic-oblong with obtusely 
cuneate, asymmetrical base, often ovate-oblong or oblong-ovate with 
rounded, asymmetrical base, attenuate and acutely acuminate at 
apex, the margin entire or slightly sinuose, 12-30 cm. long, 5-8 (-13) 
cm. broad, the acumen 1.5-2.5 cm. long, rather lustrous above, pale 
brownish when dry, glabrous, the costa and secondary nerves sub- 
filiform, prominulous, the other nerves almost unnoticeable, paler 
beneath with tawny nervation, the costa very prominent, the second- 
ary nerves about 6 pairs, prominent, ascending, decurrent, arching 
and uniting near the margin, the inferior pair forming a much more 
acute angle, the transverse tertiary nerves distant 5-10 mm. from 
470 CONTRIBUTIONS FROM THE NATIONAL 
[Figu&E 14] 
CUATRECASAS—CACAO AND ITS ALLIES 471 
each other, thin and prominent, the lesser veins forming a minute, 
prominulous reticulum; midrib, secondary and tertiary nerves gla- 
brous, those of the fourth rank subglabrous, the lesser reticulate veins 
and the areoles covered by dense, minute, whitish, sericeous tomentum 
of stellate hairs. 
Inflorescences small, axillary or extra-axillary on small branches; 
panicles short, 1-2 cm. long, branched from base, the branchlets 1-8, 
fasciculate, crowded, scarcely ramulose, densely stellate-tomentose, 
bracteolate at joints, bracteoles ovate, or ovate-lanceolate, 1-2 mm. 
long, stellate-tomentose; pedicels 2-5 mm. long, moderately thin, 
densely tomentose; sepals shortly united at base, thick, sublanceolate- 
oblong, subacute, involute at margin, ferruginous, stellate-tomentose 
outside, glabrous inside except for the minutely stellate-tomentose 
margin and glandular trichomes at base, curvate and spreading at 
an thesis, 7-9 mm. long, 2-2.5 mm. broad. 
Petal-hood thick-membranaceous, trinerved, pale and reddish 
striate, rugulose, papillose, sparsely pilose outside, oblong-obovate, 
cucullate-rounded at apex, 4 mm. long, 2 mm. broad; petal-lamina 
brownish red or rose, rotundate-subreniform, minutely crenulate, 
rugose, 2 mm, long, 2.2-2.5 mm, broad, suddenly constricted into a 
short claw, articulate at base; androecium tube 1.5-2 mm. high; 
staminodes brownish red or rose, linear-subulate, abruptly narrowed- 
acuminate at apex, the acumen often curled, densely and minutely 
pilose-muricate throughout, 5-6 mm. long, 0.7-0.8 mm. wide; fila- 
ments 1-1.2 mm. long, glabrous, arched, diantheriferous, the lobes 
of the anthers ellipsoid, 0.4 mm. long; ovary ovoid-ellipsoid, 5-ridged, 
ferruginous-tomentose, 1.5-2 mm. long, the apex whitish tomentulose; 
styles 5, thin, acute, connivent, united only at base. 
Fruits elliptic-globose, or subglobose, about 6 x 5.8 cm., rounded at 
apex, umbilicate at base, the pericarp coriaceous, minutely and densely 
tomentose, glaucous when ripe, about 5 mm. thick, the inner and 
outer layer hard, thin, the middle one carnose; seeds ovoid-oblong, 
1.4-2.1 cm. long and 0.5 x 0.9 mm. broad; pulp rather sweet, scentless, 
white; fruiting peduncle about 1.8 cm. long, 7 mm. thick. 
At the Botanische Staatssammlung in Munich there are several 
specimens from Martins' trip to Brazil which bear the annotation 
Figure 14.—a-i, Theobroma glaucum (Baker & Cope 11 and 18): A, petal from inside X 5; 
b, bud, X 2; c, pistil, X 5; D, petal laterally, X 5; e, styles, X 5; f, androecium, 
X 5; g, 2-antheriferous stamen, X 10; h, 3-antheriferous stamen, X 10; i, sepal from 
outside and inside, X 2, x-R, 7*. specioium (Archer 7619): k, petal from inside, X 5; 
L, styles, X 5; M, bud, X 2; n, petal laterally, X 5; o, androecium, X 5; p, stamen, 
X 10; q, sepal from outside and inside, X 2; r, pistil, X 5. s-y, T. velvtinum (B. W. 
1161): s, petal from inside, X 5; T, pistil, X 5; u, petal from outside, X 5; v, petal, 
laterally, X 5; w, androecium, X 5; x, stamen, X 15; y, sepal from inside and outside, 
X 2. 
080-695—64 7 
472 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
I 
. r r • 
/ if Vi ■■■ It - T Jf * ■ | - 
r
 i i' ■' ■ r * 
G ■ : 
/& 
'■ f T ' * 
f. • 
Figure 15.—a-i, Theobroma bernouillii subsp. capilliftrum (Holliday 142): a, b, c, peta 
from inside, outside, and laterally, X S; d, androecium, X S; e, stamen, X 10; f, anther; 
X 20; g, pistil, X S; h, sepal from inside and outside, X 2; I, bud, X 2. k-r, 7 
bernouillii subsp. bernouillii (Pittier 4105): k, l, u, petal from inside, outside am 
laterally, X 5; N, androecium, X 5; o, stamen, X 10; p, anther, X 20; Q, sepal fror 
inside and outside, X 2; r, pistil, X 5. s-z, T. bernouillii subsp. asclepiadiflorw 
(Wedel 1535): s, t, u, petal from inside, outside and laterally, X 5; v, androecium 
X 5; w, stamen, X 10; x, anthers, X 20; y, pistil, X 5; yy, bud, X 2; z, sepal froi 
inside and outside, X 2. 
CUATRE CABAS—CACAO AND ITS ALLIES 473 
Figure 16.—Fruits of Theobroma, X a, bernomliii subap. asclepiadiflorum (Lucas 1); 
B, bernouillii subsp, capilliferum (Cuatr. 17034); c, glaucum (Froes 20645); d, sylvestre 
(Frees 20463); e, spedosum; f, veluiinum (Benoist 516); g, nrmoraU (Cuatr. 21291). 
474 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
"Iter brasiliensis" and then the number 322 at the foot of the label; 
they are named by Martins as "T. sylvestris Martins." All the 
specimens agree with each other essentially, although some of them 
have oblong leaf blades, slightly asymmetrical or symmetrical at the 
base, and others have broad, oblong-ovate blades, very rounded and 
asymmetrical at the base; in all of them there is a longer distance 
between the basal pair of secondary nerves and the next pair than 
between the other pairs of secondary nerves; two of these specimens 
bear typical rounded, smooth fruits. One of the specimens which 
shows the number 891 on a mounting strip (Photo F. M. 19644) may 
be considered as the holotype of T. sylvestre (lectotype); it has a leafy 
branch with the basal half of a fruit, and a loose, broad, obovate- 
oblong leaf. Bernoulli, without obvious reasons, considered one of 
the specimens [890] (Photo F. M. 40709) a different species and de- 
scribed it as T. nitida Bernoulli. Another of these specimens [871] was 
named by Schumann as T. grandiflorum, and it is mixed with an 
authentic T. grandiflorum specimen on the F. M. 19641 Photograph. 
After a close examination of the series of Martins types, it was clear 
to me that there do not exist basic differences between them and T, 
spruceanum Bernoulli. Most of Martius specimens have somewhat 
thinner and smaller leaves than the current collections, variations 
which can be attributed to the habitat and to the age of the collected 
branches. 
Schumann listed T. sylvestre Mart, under his "Species dubiae/' 
The binomial Theobroma Martii was published by Schumann as a 
new name for T. nitida Bernoulli, although due to some typograph- 
ical error this name was not quoted as synonym. But Schumann 
cited "I.e. 15" and quoted unchanged the diagnosis given by Ber- 
noulli for T. nitida on page 15. Schumann considered the name a 
homonym of his T. nitidum (Poepp, & End 1.) K. Schum. (Abroma 
nitida Poepp. & Endl.). 
Theobroma sylvestre has been confused with T. speciosum, which is 
usually a larger tree, but its small axillary inflorescences, smaller, 
scentless and paler brown-reddish flowers, and its green-bluish fruit 
at maturity, distinguish it readily. 
Common names.—"Cacao azul" is generally used. Other names 
recorded are: Cacau azul, cacau!, cacauti, cacao-hd, cacao ran a, 
cacau ran a, cacau bravo. Cacao azul refers to the color of the 
mature fruit (bluish). 
Uses.—No special uses are recorded. 
Distribution.—More or less frequent along the Amazon River 
and lower part of its tributaries from Santar&n to Tonantins, east- 
ward from the mouth of the Iga. It grows on elevated ground in 
rather dry places, and appears frequently in secondary stands. 
CUATRECASAS—CACAO AND ITS ALLIES 475 
According to Ducke, it is common around Manaos, especially in 
relatively dry thickets on clay soils poor in silica. 
BRAZIL: Amazonab: "Martius Iter brasiliensis 322: Theobroma sylvestris 
Mart. Observ. 2832, Cacao sylvestris Aubl., Cacao Rana Incol. Habitat in sylvis 
ad fl. Solimoes, Prov. Rio Negro," Martina [887] (Photo F. M. 40708) (M, 
eyntype of T. sylvestris Mart.). "Martius Iter brasiliensis 322: Theobroma syl- 
vestris Mart, in si 1 vis ad Fl. Solimoes," Martius [891] (Photo F. M. 19644) 
(M, lectotype). Ibidem Martius [871] (Photo F. M, 40707) (M, syntype). 
Ibidem Martius [888] (M, syntype). Ibidem Martius [889] (M, syntype). 
"Martius Iter Brasiliensis 322, in silvis ad fl. Solimoes," Martius [890] (Photo 
F, M. 40709) (M, type of T. niiida). Manaos, Mata do Aleixo, "cacau azul," 
16 III 1945, Frdes 20556 (IAN, NY). Manaos, Colonia Jofto Alfredo, rain 
forest in noninun datable ground; small tree, flowers pale brown-rose, "cacao 
azul," 6 XII 1935, Ducke 100 (A, F, IAN, K, MG, MO, S, US). Manaos, in 
noninundatable forest; rather large tree with weak trunk, flowers brownish- 
flesh color, "cacao azul," VI 1932, Ducke 103 (F, Y). Manaos, matta, vicin- 
ity Igarap6 da Cachoeira Grande, elevated ground; small flowers on trunk and 
branches, 14 X 1912, Ducke 12187 (G, MG). Fonte BSa, matta, elevated ground; 
tree 4 m., 10 cm. diam., "cacao azul," 5 IV 1945, Fr6es 20655 (F, K, US). Ibi- 
dem, "the Indians say that it was introduced from Japura river," Frdes 20463 
(US). "Campo experimental do I.A.N,, introduced from Rio Negro, Frdes 
34949 (IAN). Mauds; small tree, flowers red, wood hard, "cacaurana," "cacau 
azul," 30 XI 46, Pires 136 (IAN). Rio Madeira, Rio Canuma, Borba munic- 
ipality; tree 4 m., elevated ground, 12 XI 1927, Frdes 33788 (IAN). Rio 
Demeni, Sumauma, Barcelos municipality; tree 8 m., fruits, flowers rose, 30 IV 
1952, Frdes 28382 (IAN). Rio Tonantins, Frdes 25554 (IAN). 
Map S.—Geographical distribution of Theobroma speciosum # and 7*. glaucum Q 
476 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
ParA: "In vicinibus Obidos, Prov. Para, Dec. 1849," Spruce 166 (Photo 
F. M. 40702) (M, holotype and lectotype of Theobroma spruceanum Bernoulli; 
isotypes BM, WU). Obidos, "cacao azul," 7 I 1904, Ducke 4878 (BM, G, MG, 
P, US); ibidem, matta, 11 V 1905, Ducke 7216 (MG). Ibidem, rain forest, ele- 
vated ground; small tree, fruit green glaucous when ripe, flowers pale brownish 
reddish, "cacao-azul," 10 I 1920, Ducke 14734 (S, U), Sao Jorge, municipality 
of Faro; small tree in rain forest, flower dark red on branches, "cacau azul," 
11 XI 1950, Black & Ledoux 50-10644 (IAN, UC, U). OriximinA, Las Trombetas, 
flowers on trunk and branches, ripe fruit green, "cacao azul," "cacao-hu," 8 XII 
1906, Ducke 7822 (BM, G, MG, US). Lazo de Faro, above Tanaenera, forest on 
elevated ground, "cacau azul," 12 II 1910. Ducke 10669 (MG). Alenquer, 
Estrada da Vila do Curud, municipality of Obidos, sandy ground with vegetation 
of Bertholletia and Attalea; tree 2.5 m. in rain forest, 4 III 1953, Frdes & Filho 
29465 (IAN, U). "Km 23 da BR-17 Est. a direita," firm land, sandy, natural 
forest; flowers yellowish red, edible fruits, 12 X 1955, "cacao bravo," INPA 
(.Dionisio) 2125 (IAN), 
3. Theobroma speciosum Willd. 
Figures 5, 12, 13,14,16,18; Map 5; Plates 2, 3 
Theobroma speciosum Willd. ex Spreng. Syst. Veg. 3:332. 1826; Bernoulli 
(1869) 8, pi. 3, fig. $; Schumann in Mart. (1886) 74; Jumelle (1899) 30, 
fig. 16; De Wildeman (1902) 95; Huber (1906a) 273; Ducke (1925) 130; 
(1940) 270, pi 1, fig. S, pi. 2; (1954) 13; Addison & Tavares (1951), pi. 5, 
fig. 8, pi. 6, fig. B, pi. IS, fig. 6; Guatrecasas (1956) 658. 
Theobroma quinquenervia Bernoulli, Uebers. Art. Theobroma 8, pi. 3, fig. 3. 
1869. 
Theobroma speciosum var. quinquenervia (Bernoulli) Schum. in Mart. Fl. 
Bras, 12(3):75. 1886: Jumelle (1899) 32, fig. 16; De Wildeman (1902) 95. 
Theobroma speciosum var. coriaceum Huber, Bol. Mus. Goeldi 4:586. 1906. 
Theobroma guianensis sensu Chevalier, Rev. Inter. Bot. Appl. 26:274, 1946; 
Le6n (1960) 318, 319, fig., pro parte, non Gmelin. 
Sapokaia brasiliensia Rich, ex Chevalier. Rev. Int. Bot. Appl. 26:275. 1946, 
as synonym. 
Types.—Siber: " Hojfmannsegg" in Herbarium Willdenow no. 
3680 (B). Spruce 1737, Brazil, Barra do Rio Negro (of T. quinque- 
nervia). Huber 1567, Peru, Ucayali (of var. coriaceum). 
Tree up to 15 mm. tall; trunk about 20 cm, in diameter with light 
gray, smooth bark; sympodial growth by lateral, sub terminal upright 
shoots; primary branches tern ate, usually furcate, spreading; branch- 
lets terete, smooth, more or less pulverulently stellate-pilose, later 
glabrate, grayish brown; crown rather narrow; stipules subulate, 
short, stellate-tomentosc, soon deciduous. 
Leaves firmly coriaceous, distichous; petiole robust, sub terete, 
densely stellate-tomentulose, 8-14 (-20) mm. long, 2-4 mm. thick; 
blades usually large, ovate-oblong or elliptic-oblong, broadly rounded 
or very obtuse and asymmetrical at base, exceptionally (in ortho- 
tropic branches) cuneate and symmetrical, attenuate near the apex, 
abruptly and acutely acuminate, the margin entire or very slightly 
sinuose, often slightly revolute, 20-40 cm, long, 7-18 cm. broad, the 
CUATRECASAS—CACAO AND ITS ALLIES 
477 
Figure 17.—Leaves of Tkeobrotna, X : a, glaucum, from lateral branches (Hartley 
& Holl. 74); b, glaucum, from orthotropic branches (Baker 37); c, bernouiltii subsp. 
asclepiadifloTum (Wedel 681); d, hernomllu subsp, bernouiltii (Pittier 3199); e, bemouillii 
subsp. captlliferum (Cuatr. 16160). 
478 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
acumen 1.5-2.5 cm. long; lustrous above, green, usually pale brown or 
olivaceous when dry, glabrous, or with a few stellate hairs on the 
nerves, the costa and main nerves filiform, conspicuous, the minor 
veins less noticeable or obsolete, pale cinereous beneath with tawny 
nervation, the costa thick, very prominent, the 6-8 pairs of secondary 
nerves prominent, subcurvate ascending, near the margin thinner, 
curving, decurrent and anastomosing, the inferior pair stronger, 
forming together with the midrib a trinerved base, the transverse 
tertiary nerves thin but prominent, 5-15 mm, distant from each 
other, sometimes the basal more conspicuous, forming together with 
main nerves a basally 4- or 5-nerved leaf; minor veins prominulous, 
reticulate; main nerves glabrous or subglabrous with sparse, stellate 
hairs and scattered callose dots, the tertiary glabrous or subglabrous, 
the minor reticulate veins and the areoles covered by dense minute 
whitish sericeous tomentum of stellate hairs. 
Inflorescences on trunk, forming delicate panicles gathered in many- 
flowered bunches borne on woody, short, tuberculose branches, 
often very showy with up to 250 dark-red or purplish-red fragrant 
flowers; panicles 3-10 cm. long, dichotomously furcate-branched from 
the base, the branches thin but rigid, reddish and covered with a 
minute, whitish tomentum, the terminal branchlets (peduncles) thin, 
flexuous, 10-25 mm. long, 3 bracteolate and articulate to the pedicel 
at apex; pedicels slender, 5-20 mm. long, minutely tomentulose; 
bracteoles minute, linear or linear-triangular, 1-2 mm. long, 0.2- 
0.5 mm. wide, very soon deciduous; buds ovoid, 7-9 mm. high, rather 
reddish, with 5 longitudinal, white-tomentose, prominulous com- 
missural lines, sparsely stellate-pilose; sepals rather thick, oblong, 
subobovate-oblong, attenuate toward the base, abruptly narrowed 
and subobtuse at apex, the margin incurved with a minutely, whitish 
tomentulose strip inside, the apex shortly cucullate-inflexed, the inside 
purplish and glabrous, except for glandular trichomes at base, the 
outside sparingly stellate-pilose, usually one free and the others 
united one third or almost completely by pairs, 10-12 mm. long, 
3.5Ht mm. broad; petal-hoods thick-membranaceous, trinervate, 
whitish with red lines, oblong-obovoid, attenuate-clawed at base, 
rounded cucullate at apex, with sparse, thin, spreading hairs outside, 
6-7 mm. long, about 4 mm. broad; petal-lamina rather thick, red or 
dark red, transversely elliptic, subtruncate, slightly emarginate and 
mucronulate at apex, abruptly cuneate-attenuate at base, entire or 
slightly erose at margin, conspicuously (especially by transmitted 
light) reticulate-veined, 5-7.5 mm. long, 7-9 mm. broad; androecium 
tube thick, about 2-3.5 mm. high, sparsely stellate-pilose; staminodes 
purplish red, subulate, thick, the apex thinner and curled, minutely 
muricate-pilose, 5-7 mm. long, 1.2-1.8 mm. wide; filaments glabrous, 
CUATRECASAS—CACAO AND ITS ALLIES 479 
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Figure 18r—Indument on the underside of leaf in: a> Theobroma bicolor (Killip & Smith 
30006); b, 7\ sylvestre (Ducke 100); c, T. speciosum (Ule 9629); d, T. vtlutinum (Benoist 
516). A, B, and C X 30, D X 20, 
480 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
about 2 iniii. long, curved, dilated at base, minutely 3-furcate at apex, 
triantheriferous; anther lobes ellipsoid, about 0.4 mm. long; ovary 
ovoid-ellipsoid, 2-3 mm. long, 5-ridged, whitish velvety-tomentose; 
styles 5, subfiliform, 1.2-2 min. long, glabrous, eonnivent, only united 
at base. 
Fruit globose-ellipsoid, about 10 cm. long and 7-8 cm. broad, almost 
smooth, with 5 more or less conspicuous (when dry depressed) costac, 
shortly and densely torn en tose-velvety, yellow when ripe; pericarp 
about 5-6 mm. thick, the inner layer coriaceous, smooth, very hard, 
about 0.5-1 mm. thick, the middle tissue about 3—4 mm., carnose, the 
outer layer coriaceous but less hard than the innermost and becoming 
rugose after the shrinking of the intermediate layer by drying; seeds 
about 20-26, surrounded by whitish, sweet, scentless pulp, ovoid- 
oblong or ellipsoid-oblong, 24-26 mm. long, 13-14 mm. broad, 10- 
12 mm. thick, the episperm thick (about 1 mm.), coriaceous with the 
middle layer becoming gelatinous; embryo white, oblong, covered by 
a very thick pellicle, 22-24 mm. long, 10-11.5 mm. broad, 9-10 mm. 
thick; germination epigeous. 
The leaves of the type of T. speciosum are Iong-petiolate, cuneate 
and trinerved at the base. Theobroma quinquenervium Bernoulli was 
described from a specimen with short-petiolate, broadly oblong 
leaves, asymmetrically rounded at the base. In the first type the 
margins of the leaves are close and parallel to the prominent, basal 
pair of secondary nerves; in the second the margins of the leaves are 
broadened and remote from the prominent basal pair of nerves, and 
an additional lower tertiary nerve on each side makes the base of the 
blade somewhat 5-nerved. The latter type of blade, borne on a short, 
stout petiole, is the common one in the species. Leaves with slender, 
long petioles, thickened at both ends, and cuneate blades are seldom 
found; they appear on young, orthotropic terminal branches. This 
dimorphism was already noticed by Huber who first united T. quin- 
quenervium and T. speciosum. 
The type specimen was collected near Bel6m de Par& by Siber who 
was sent on a collecting trip to Brazil by Hoffmannscgg; it is pre- 
served in the Berlin-Dahlem Botanical Museum in the Willdenow 
Herbarium. I have been able to study this specimen thanks to the 
kindness of Prof. Werdermann and Prof. Melchior, The photo- 
graph F.M. 9640 is from a specimen at B now destroyed, which agrees 
perfectly with the type in the Willdenow Herbarium, of which it was 
undoubtedly a duplicate. In his monograph, Chevalier made this 
species a synonym of T. guianense (Aubl.) Gmel., but this is a con- 
fused species, the identity of which is discussed in this paper. 
Spruce and Ducko called attention to beauty of this tree in blossom, 
according to Spruce "one of the prettiest things I have seen." 
CUATRE C ASAS—CACAO AND ITS ALLIES 481 
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Figure 19.—a, b, Detail of nervation at the underside of the leaf in: a, Tktobroma speciosum 
(Ule 9609); b, T. cacao (Cuatr. 7756). c, d, indument on the underside of the leaves 
in: c, T. glaucum (Bart. & Holl. 74); d, T. bernouilliisubsp. capilliferum (Cuatr. 16160). 
A and B X 2, C and D X 25. 
482 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Theobroma speciosum can grow to be 15 m. high, with a few whorls 
of dichotomous leafy branches near the top of a long branchless stem, 
which can bear abundant, large, caulinc inflorescences. These may 
form large cushions of showy, blood-red, wine-red, or purplish-red 
flowers which give off an intense lemon- or orange-skin odor (Ducke). 
The outer pulp of the seeds is sweet but scentless. It is said to be easy 
to grow in gardens. 
Common names,—Cacauf, cacauu. Other recorded names or 
different ways of spelling are: cacao-y, cacao-u, cacau, cacaohy, 
cacau-i, cacao-i, cacaofllo, cacau-rana, cacao-rana, cacao biaro, 
cupuy, cupuyh, cacao do matta, cupurana, cacao azedo, cacao sacha 
(Peru), chocolatillo (Bolivia). 
Uses.—The pulp is eaten by natives. The seeds are used very 
occasionally to prepare low quality chocolate. 
Distribution.—This species extends throughout the Amazonian 
Hylaea except in the northwestern section, from the state of MaranhSo 
at Cururupu to Acre and Madre do Dios in eastern Bolivia and the 
Ucayali River (Loreto) in Peru. It grows on noninundatable ground 
in rain forests as well as in not too humid places and it appears also in 
secondary growth near towns, but it is never of frequent occurrence. 
BRAZIL: "Brcsil. Herb-Lusit," "Herrania paraensis," "Theobroma subin- 
canum Mart.?/' Geoffroy [St. Hilaire?] s.n. (P), "Polyadelphia Decandria 
Theobroma speciosa foliis acuminatis integerrimis subtus toraentosis. Floras 
purpurei. Habitat in Para Brasilia," "Theobroma speciosa (W.) Spreng. Bernoulli 
determ." Hoffmannsegg, W. (B, Herb. Willdenow 3680, holotypus). "Ex herb. 
Linn. Theobroma speciosa Brasil Col." Siber (B), specimen destroyed (photo- 
graph F. M. no. 9640); identical with the type in the Willdenow Herbarium. 
ParA: Bel£m, noninundatable forest near city; small tree with dark-red 
flowers on trunk, "cacaohy," IX 1936, Ducke 281 (A, F, K, MO, NY, S, US). 
Bel6m, Jardim BotAnico do Museo Goeldi; slender tree, inflorescence on trunk, 
petals dark red, sepals reddish stained, fruit 5-parted, yellow when ripe, "cacau-i," 
26 VIII 1942, Archer 7619 (IAN, K, NY, USD A). Ibidem, slender tree, 
"cacao-i," 29 X 1942, Archer 7721 (IAN, USD A). Ibidem, small tree, flowers 
deep crimson, calyx light pink, leaves deep green above, gray green beneath, 
cultivated, 15 VII 1946, Schultcs & Silva 8066 (GH). Ibidem, 19 X 1945, Pires 
& Black 695 (IAN), 740 (IAN). Braganga, "cacao claro," XII 1899, Huber 
1748 (G, MG). Tapcrinha, at Santarem, secondary forest on elevated land; 
small tree, red flowers on trunk, "cupuy," Gimberger 802 (WU). Rio Curuauna, 
Cachoeira do Portao, region of Planalto de Santardm, rain forest, XI 1954, Fries 
31414 (IAN). Santardm, hills, "cacao do Mat to," Jobert 903 (P), Near Obidos, 
Prov. Par&, XII 1849, Spruce s.n. (K, Herbarium Hookcrianum 1867). Forest 
near Obidos, XII 1845; slender tree 40-50 ft., flowers on the naked stem, leaves 
on the top of the tree only, "cacao-rana," Spruce 456 (K, Herbarium Benthami- 
anum, 1854, P, BM). Monte Alegre, region of Colonia da Mulata, elevated 
ground; tree 8m., red flowers, 28 IX 1953, Frdes 30432 (IAN). Breves, VII 
1956, Pires, Frdes, & Silva 5886 (IAN). Belterra, capoeira, way to Pindobal; 
tree 7 m.f red flowers on trunk with lemon scent, leaves white beneath, 31 XI 1947, 
Black 47-1889 (IAN, NY). Region Tapaj6s, B5a Vista firm land; tree 4 m., 
CUATRECASAS—CACAO AND ITS ALLIES 483 
10 cm, [diam.], flower color wine, fruits on the trunk, eaten by Indians, "cupuhy," 
30 VIII 1932, Capucko 397 (IAN, F). Tapaj6s, Vila Braga; tree, dark-red 
flowers borne on the trunk, 27 X 1908, Snethlage 10044/b (MG). Upper Cupary 
River, plateau between the Xingu and Tapaj6s Rivers; tree 25 ft. high, 2 inches 
in diam. breast high, in high forest, flowers red on old wood, "cacaorana," IX 
1931, Krukoff 1117 (A, G, K, NY, P). Ibidem; tree near river shore; flowers 
dark red, borne on large branchlets; wood used for "f&rinha" containers (to keep 
it dry), "cacaorana," Krukoff 1080 (A, BM, G, K, MICH, MO, NY, S, U). Rio 
Cumina-mirim, forests at NE; ripe fruit yellow, only on trunk, "cacaorana," 
16 XII 1906, Ducke 7975 (MG). Oriximina, Las Trombetas, "cacao rana," 
flowers only on trunk, 8 XII 1906, Ducke 7884 (G, MG). 
Amazon as: Near Barra, Prov. Rio Negro, Aug. 1851; forests south of Rio Negro; 
tree 20 feet, straight with a whorl of branches only at summit, which are twice 
or thrice dichot[omous]; trunk almost completely clad with flowers, which have 
fine odor of bruised orange leaves; petals cucullate, claw pink, white limb dull 
crimson with dark veins. The subul[ate] processes blood red; one of the prettiest 
things I have seen, Spruce 1737 type of Theobroma quinquenervium Bernoulli 
(M, holotype, Photo F, M. 40703; isotypes: BM, E, F, G, GH, GOET, K, LD, 
LE, OXF, P, WU) (Photo F. M. 9639 from Berlin), Rio Marmellos, near 
mouth; flower deep salmon red, buds dark crimson, bark smooth, light gray 
white, leaves pale dull beneath, dark green glossy above, branches umbelli- 
form at top, trunk 6-7 inches in diameter at base, 20-22 feet tall, inflorescences 
200-flowered; seeds give low grade chocolate, 2-12 VIII 1945, "cacao azedo," 
SckuU.es tfc Cordeiro 6507 (AMES, F, IAN). Amazonas, Rio Capitare, munici- 
pality of Codajas, elevated ground, high forest; tree 8 m,, red flowers, 3 IX 1950, 
Fr6es 26526 (IAN). Manaos, Estrada do Aleixo; tree 5 m., fruits on trunk, 
14 V 1953, Frdes 30180 (IAN). 
Guapoke: Porto Velho, Estrada de Rodagem, Km 8, Viana, disturbed 
forest, elevated land; small tree, red flowers with lemon scent, 31 V 1952, Black, 
Cordeiro, & Francisco 52-14655 (IAN). 
Goias; Margen do Rio Araguaia; tree 4 m., red flowers, 13 VI 1953, Fries 
29732. 
Ache (Territorio del): Rio Acre, Seringal San Francisco; tree 5-20 m., 
black-purplish flowers on stem, VII 1911, Ule 9609 (G, K, L), Vie 14448 (MG), 
Near mouth of Rio Macauhan (tributary of Rio Yaco), Lat. 9°20' S., Long, 69° W., 
on firm land; tree 40 feet high, 5 VIII 1933, Krukoff 5295 (A, F, K, LE, M, MICH, 
MO, NY, S, U, UC, US). 
Rio de Janeiro: Quinta de Sao Christovas, 10 I 1876, Glaziou 9633 (C, P). 
Jar dim Bot&nico, flowers on trunk and main branches, V 1944, Camargo 2395 
(IAN). 
PERU: Loreto: Region of the middle Ucayali, rain forest, Yarina Cocha, 
155 m. alt., elevated ground; tree 12-15 m.t 35 cm. thick, the first branch at 
7 m. from ground, flowers with strong anise scent, sepals red crimson, petals dark 
crimson-striped on white ground, stamens less crimson than petals, pistil whitish, 
"cacao saeha," 22 IX 1925, Tessmann 5398 (G, M, S). Rio Ucayali, Paca, 21 
VII1898, HvJber 1567 (Holotype of var. coriaceum Huber, MG; isotypes BM, US), 
(photo F, M. No. 1567). 40 km. south of Pucallpa, rain forest of the Amazon basin 
virgin rain forest on loamy sand, Ellenberg 2565 (U). 
BOLIVIA: Km. 7 on road Guayaramerin-Cochuela Esperanza, [Prov. Vaca 
Diez, Depto. Beni], 120 m. alt., "chocolatillo"; growing wild in rain forest; flowers 
wine red, borne on cushions along the whole, rather slender trunk, dichotomous 
branches slender, pendent, fruits small. I have seen also a specimen in Brasilia, 
where it is called "cacau-i"; 9 IX 1954, Patino s.n. (GH). Junction of Rivero 
484 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Beni and Madre do l)ios; pulp edible and equal to that of T. cacao, seeds white, 
not used, VIII 1886, Rwby 654 (BM, E, F, G, GH, K, LE, MICH, MO, NY, P, 
US). Unduavi, 10,000 ft., Rmby 647 (US). 
4. Theobroma velutlnum Benoist, Bull. Mus. Hist. Nat. Paris 27: 113. 1921. 
Figures 3, 12, 13, 14, 16, 18; MAP 1. 
Herrania guianensis Sagot ex Schum. in Mart. Fl. Bras. 12(3): 75. 1886. 
Theobroma speciosum sensu Uittien in Pulle, Fl. Surinam 3:45. 1932. 
Theobroma sp. 4, Uittien in Pulle, Fl. Surinam 3:46. 1932. 
Herrania guyanensis Sagot in Chevalier, Rev. Int. Hot. Appl. 26:275. 1946 
(as synonym). 
Type.—Benoist 516, French Guiana. 
Branches terete, subcinereous, densely appressed stcllatc-tomentose 
or becoming glabrate, grayish brownish; branchlets densely toraentose. 
Leaves large, firmly coriaceous; petiole robust, sub terete, densely 
appressed cinereous-tomentose, when old transversely rimose, 12-14 
mm. long; lamina oblong-ovate or ovate-oblong, broadly rounded and 
more or less asymmetrical at base, suddenly narrowed and long- 
acuininate at apex, the margin slightly revolute, entire or very slightly 
sinuosc, 28-40 cm. long, 16-21 cm. broad, including the acumen, this 
about 3 cm. long, pale olivaceous above (when dry), subnitidous, 
slightly and broadly bullate, glabrous except for the midrib, this 
sparsely pilose towards the base, the costa and secondary nerves thin, 
the tertiary nerves filiform, the minor veins less apparent or obsolete, 
softly velutinous beneath, ochraceous-cinereous, the midrib very 
prominent, the 5-7 pairs of secondary nerves prominent, ascending, 
curved near the margin and anastomosing, the longer pair stronger, 
more separated from the next, the transverse tertiary nerves promi- 
nent, others broadly reticulate, prominulous, the lesser veins forming a 
minute, prominulous reticulum, the minor reticulum and its areoles 
densely tomentose with minute, white, dense, subappressed, stellate 
hairs, the other nerves and the sides of the midrib covered with abun- 
dant, long, delicate, stellate hairs with long, thin, spreading rays. 
Inflorescences many-flowered, borne on lignose, tuberculate branches 
on the trunk, the panicles fasciculate, ramose from the bases, 5-12 cm. 
long, the branchlets moderately thin, cinereous, hirtellous-tomentel- 
lous with thin, mediocre, stellate hairs, the terminal (peduncles) 
thinner with 2 or 3 deciduous bracteoles at the end; pedicels slender, 
6-18 mm. long; bracteoles at the joints minutely lanceolate, hirtellous, 
deciduous, 1—1.5 mm. long; sepals thick-membranaceous, spreading, 
shortly united at base, lanceolate-oblong, subacute, glandular at base, 
otherwise glabrous inside, with fine stellate hairs copious on the 
outside, the margin minutely tomentulose, about 10 mm. long and 
3.5-4.5 mm. broad, usually one free, the others united by pairs. 
Petal-hoods 6-7.2 mm. long, about 3 mm. broad, membranaceous, 
obovate-oblong, cucullate-rounded at apex, 3-nervate, the median 
CUATRECASAS—CACAO AND ITS ALLIES 485 
vein furcate, outside sparsely, weakly pilose, the end auriculate- 
emarginate, articulate to the erect lamina, 6-7.2 mm. long, about 
3 mm. wide; petal-lamina sessile, subtrapezoid, sub truncate or often 
slightly sinuate or 3-toothed at apex, abruptly narrowed into a short 
claw at base, moderately thick, but venation conspicuous by trans- 
mitted light, glabrous, somewhat rugulose-papillose toward the base, 
5 mm. long, 6-6.5 mm. broad; androccium-tube about 2 mm. long and 
2.4 mm. broad, minutely papillose-pilose; staminodes purplish red, 
lanceolate-subulate, thick, suddenly narrowed into a short, crisp, 
acute acumen at apex, minutely muricate-pilose, about 6 mm. long 
and 1.5 mm. wide; filaments triantheriferous, 2.5 mm. long; ovary 
ovoid-oblong, about 2.2 mm. long, 5-ridged, tomentose; styles 5, 
connivent, adherent, easily separable, glabrous, 2 mm. long. 
Fruits ellipsoid, densely and softly stellate-pilose-velutinous, 5- 
eostate, the ribs thick, very prominent, the surface smooth, 8.2-9 cm. 
long, 6-6.3 cm. broad, the pericarp 3-4 mm, thick; seeds usually 25-30 
in each pod. 
This species is closely related to T. spedosum, having very similar 
flowers and leaf outline. However, T. velutinum is very different on 
account of the structure of the fruits and the indument. The ellipsoid, 
densely velvety fruits have 5 longitudinal, very prominent, typical 
ribs, a character only known in this species. The leaves beneath have 
a soft, velvety indument of long, thin, stellate or dendroid, more or less 
densely distributed hairs on the whole nervation and a lower layer of a 
short, dense tomentum of minute, stellate, intricate hairs covering 
the areoles and the minor reticulate veins. Also, the terminal 
branches are densely tomentose and the inflorescence branchlets and 
pedicels have copious, rather long, fine hairs. 
Theobroma velutinum is only known from French Guiana and the 
neighboring Dutch side of Maroni River valley. The excellent foliage 
and fruiting collections made by Benoist and recent other collections 
by members of French and Dutch Forest Services, especially the 
flowering specimens B. B. S. 1161, have facilitated a complete descrip- 
tion of the species. Its inflorescences are cauline, many-flowered, 
and often showy, like those of T. spedosum. The Sagot collection 
1206, preserved in several herbaria and consisting only of large 
inflorescences, belongs to this species. Sagot named it Herrania 
guianensis and left an accurate unpublished description which is 
attached to the specimens in Paris. 
Common names.—Bouchi-cacao, cacao sauvage, cacao. 
Uses.- Reported to yield edible seeds comparable to cacao but no 
information is given on the quality of the product. 
Distribution.—French and Dutch Guiana in the valley of Maroni 
River. 
486 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
SURINAM: 4X 1950, B. W. 1161 (U). Placer l'Arver, 31 X 1918, Gonggryp 
4108 (U). Flur of Marowyne, Reu naar L. etwa en Tupanahoni, No. 47 Maro- 
wyne, 26 XI 1918, Gonggryp 4127 (U). Flur Tapanahoni, III 1922, No. 33 
Tapanahoni, Gonggryp 4148 (U). 
FRENCH GUIANA: A 1 kilometre plus ou moins du camp de transports de 
Charvein, le long de cherain qui conduit a la leproserie de TAcarouany, 8 I 1914, 
Benoist 516 (P, holotype, foliage and fruits). Crique Serpent, rive droit & 1 
kilometre de la crique terrain en pente, & proximity immediate d un ravin rocheuse; 
arbuste ayant 6t6 rep&6 au moment de sa fructification en fdvrier 1951; abattu 
depuis et pourvu de re jets de 3 m. de haut; produit de gousses cdt^es contenant 
des grains comestibles, "bouchi cacao" (Paramaka), 12 XII 1952, BAFOG 136M 
(P). Placean No. 2, Carreau No. 3, route de mana, terrain sablonneaux; fruits 
jaun&tres, gousses ovoides de 7 & 8 x 10 & 12 cms. d<5hiscents, sparses sur le tronc 
de l'arbre, "bouchi cacao," "cacao sauvage," 19 III 1956, BAFOG 7386 (P). 
Karouany, 1858, "Flores atropurpurei suaveolentes, fructus o vat us, pentagonus, 
breviter tomentosus cacao sativa paulo minor, folia non vidi, flores e ligno 
prodeunter, "cacao," "Herrania guianensis Sagot," 1858, Sagot 1206 (BM, G, 
K, P, U, WU); type of Herrania guianensis Sagot (only inflorescences and flowers 
in all the specimens; a long and accurate description made "in vivo" by Sagot 
is attached to the specimens at Paris). 
5. The*ibroma glaucum Karst. 
Figubes 2, 10, 14, 16, 17, 19; MAP 5; PLATE 4 
Theobroma glaucum Karst., Linnaea 28:447. 1856; Triana & Planch. (1862); 
Bernoulli (1869) 10; Jumelle (1899) 34; De Wilde man (1902) 97; Chevalier 
(1946) 277. 
Theobroma cahdesmis Diels, Notizbl. Bot. Gart. Berlin 14:336. 1939; Baker, 
Cope & al. (1954), Jigs. 10, 12; Cuatrecasas (1956) 655; Lefin (1960) 314, 
317, fig. 
Types: Karsten s.n. Colombia, San Martin. SchuUze-Rkonhof 
2312, Ecuador, Papayacu (of T. calodesmis, formerly in Berlin). 
Tree 8-15 m. high; stem up to 30 cm. in diameter, with grayish, 
inside reddish bark and white wood; sympodial growth by lateral, 
sub terminal, upright shoots; primary branches ternate, regularly 
dichotomous, spreading, deciduous when old, the terminal minutely 
stellate-pulverulent with additional simple, spreading hairs, soon 
glabrate, smooth, rather shining, dark brown or somewhat purplish; 
stipules linear-subulate, 4-5 mm. long, sparsely stellate-pilose, soon 
deciduous. 
Leaves coriaceous, rather rigid, distichous; petiole robust, densely 
ferruginous or brownish tomentose with stellate hairs, transversely 
rimose when dry, 0.8-1.8 cm. long; blades oblong-ovate or ovate- 
oblong, broad in the lower third, obtusely cuneate at base, narrowed 
near the apex, prolonged with a long slender appendage, the margin 
entire or slightly sinuate and slightly revolute, 16-36 cm. long, 7-13 
cm. broad, the acumen 2-3.5 mm. long, shining above, green, pale 
olivaceous brown when dry, apparently glabrous but with sparse 
mediocre stellate hairs and callose scar-dots on the nerves, the costs 
and secondary nerves prominently filiform, the others slender, more or 
CUATRECASAS—CACAO AND ITS ALLIES 487 
less noticeable, somewhat cinereous beneath, glaucous or pale rosy, 
with a glabrous aspect but the rather shining, pale brownish principal 
nerves sparsely callose-dotted and with very scarce ferruginous, 
stellate hairs, the small veins glabrous, the areoles covered with a 
very appressed microscopic tomentum of minute, white, stellate hairs, 
the costa very prominent, the 4 or 5 pairs of secondary nerves promi- 
nent, the basal one at an acute angle (remote from the margin), ascend- 
ing, the others curved-ascending, near the margin becoming slender, 
decurrent, curving, anastomosing, the cross-tertiary nerves thinner, 
prominent, 3-10 mm. distant from each other, the lesser veins 
minutely prominulous-reticulate; leaves of the orthotropic spreading- 
puberulous branches long-petiolate, with the blades attenuate- 
cuneate at base, the lower pair of nerves very close to the margin, 
the petiole slender, 3.5-4 cm. long. 
Inflorescences on the trunk, often many-flowered and showy, with 
up to 200 flowers, the base woody-tuberculate; branches 3-many, me- 
diocre, 4-6 cm. long, furcate-ramose from near the base, the branchlets 
fastigiate, angulate, rather rigid, ferruginous-tomentose, the terminal 
(peduncles) moderately robust, 3^4 mm. long, articulate to pedicel, 
this 5-15 mm. long, striolate, slightly thicker, tomentellous, the 
subtending bracteoles minute, ovate-lanceolate, about 1 mm. long, 
very soon deciduous; buds ovoid, round at base, subacute at top, 
densely stellate-tomentose, 8-9 mm. long, about 6 mm. broad. 
Calyx umbilicate; sepals thick, lanceolate-oblong, acute, inflexed 
at apex, connate for 2 mm. at base, densely and appressed stellate- 
tomentose outside, within minutely, whitish stellate-pilose near the 
margin and glandular at base, otherwise subglabrous, 12-13 mm. long, 
3.5-4 mm. broad, curved-spreading after anthesis. 
Petal-hoods light red, oblong-obovate, shortly unguiculate at base, 
rounded cucullate at apex, the end emarginate, biauriculate, articulate 
to the lamina, the 3-nerves prominent inside, thin, with spreading, 
weak, sparse hairs outside, 5-6 mm. long, 2.5-3 mm. broad; petal- 
lamina red crimson, thick, minutely rugose, and more or less trans- 
lucid-venulose, glabrous, suborbicular or broadly elliptic, subsessile 
at base, abruptly contracted into a short claw, minutely sinuate at 
margin, 5.5-7 mm. long, 5-6.5 mm. broad, the claw 0.5 mm. long. 
Staminal tube about 2 mm. high and 2.5 mm. in diameter; stami- 
nodes red crimson, erect, lanceolate-subulate, acute at apex, fleshy, 
minutely muricate-pilose, 10-12 mm. long, 1.4-1.8 mm. wide above 
base; filaments flexuose, 2,5-3 mm. long, glabrous, shortly 2 or 3 
furcate at apex, bearing 2 or 3 anthers, the loculi ellipsoid, convergent, 
0.5-0.6 mm. long; ovary oblong, about 2 mm. long, 5-ridged and sul- 
cate, hirsute-tomentose; styles connivent, 3 mm. long, united only 
at base. 
680-695—64 8 
488 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Fruits ellipsoid-oblong, obtusely pentagonal, broad and umbilicate 
at base, more or less attenuate and subacute at apex, the pericarp 
1 cm. thick, coriaceous, rigid, densely and minutely velutinous- 
tomentose, bluish greenish, 11-13 cm. long, 5.5-9 cm. broad; seeds 
2-2.3 x 1.2-1.4 x 0.9 cm.; fruiting peduncle robust, 7-8 cm. long, 1-1.5 
cm. thick; germination epigeous. 
According to Baker and his associates the cotyledons are white and 
the pulp is pale orange and of a very sweet taste. 
The type collection of T. calodesmis was destroyed during the war 
in Berlin; it was collected by Hertha Schultze-Rhonhof near Papayacu 
at about 200 m. altitude on the Bonanza River, a tributary of the 
Pastaza River in eastern Ecuador. The description given by Diels 
makes it possible to identify his species perfectly with several Amazo- 
nian collections from nearby regions of Colombia, Peru, and Brazil. 
The sterile type specimen of Theobroma glaucum, collected on the 
Llanos de San Martin, agrees perfectly with a specimen that is almost 
a topotype, collected by Philipson, Idrobo, and Fern&ndez in the 
foothills of the Sierra Macarena, Intendencia del Meta. I have no 
doubt that all these collections represent the same species, which 
extends from the upper Orinocia to upper Amazon basin on both sides 
of the great river. 
Diels did not see fruits but gave an accurate description of the 
foliage and flowers; he related his species to T. speciosum on account 
of the texture and to men tu m of the leaves, and also to T. bernouillii, 
but he says that the leaves are broader, the inflorescences larger, the 
flowers larger, and the staminodes longer. 
Common names.—Cacao de monte, cacao silvestre, chucu (Rio 
Papayacu [Schultze-Ronhof]), "bicco" Rio Apaporis). 
Uses.—According to Karsten, the seeds are used as cacao by the 
natives, being very similar to the true cacao. Schultze-Rhonhof gives 
the indigenous name "chucti" for the fruit which, according to her, 
is very much appreciated by the natives. 
Distribution.—This species grows in the upper Amazon basin 
(northwestern section) along the rivers Caqueta, Cagu&n, Putumayo, 
Vaup4s, Guainia, Inirida, and Apaporis in Colombia and along the 
Colombian boundary with Ecuador, Peru, and in western Brazilian 
Amazonas; at the northern end of its range it enters the Orinoco basin 
into the Meta drainage in Colombia. 
It grows in the humid rain forests from the lower level of the great 
rivers to an altitude of 450 m. in forested hills. 
COLOMBIA: Meta: Villaviconpio, Llanos de San Martin, Karsten, s.n. (WU, 
holntype) (photo P. M. U2205). Sierra Macarena, Cano Ycrly, 450 m., dense, 
humid forest; unbranched tr«e 10 in. high, bundles of cauli Morons fruits grern, 
24 XI 1949, Philipson, Idrobo, & Fern&ndez 1552 (BM, COL, US). 
CUATRECASAS—CACAO AND ITS ALLIES 489 
Putumayo: Vicinity of Mocoa; tree 6-7 m., growing fully exposed in meadow, 
trunk 23 cm. thick at base, branched inflorescence (dead) borne on trunk and 
branches, pod broadly pentagonal with 5 very shallow furrows, 13 cm. long, 
9 cm. diam., blue green in colour, fruit pedicel S cm. long, 1.5 cm. diam., with 
abscission ring 1.5 cm. from pod base, in colour pale green with fine white hairs, 
17 III 1953, Holliday & Cope T/79 (COL, TRIN, US). Ibidem; sterile tree 15 m. 
in forest, undoubtedly similar to T/79, shoots from below jorquette, Holliday 
& Cope T/79A (COL, TRIN, US). Rfo Legui'zamo, Laguna Primavera, 3 IV 
1953; tall tree 15 m., obviously cauliflorous, Holliday & Cope T/94 (COL, TRIN, 
US). Rfo Legufzamo; tree 16 m., with old trunk inflorescences, no flowers or 
fruits, 5 IV 1953, Holliday & Cope T/96 (COL, TRIN, US). 
CaquetA: Rio Cagudn, Camp 4; branched tree 10 m., found in good flowering 
condition, flowers in large inflorescences on trunk, 27 IV 1953, Holliday & Cope 
T/118 (COL, TRIN, U, US). Ibidem; tree 15 m., branched, two immature pods, 
on sloping land near T/114, 26 IV 1953, Holliday <£ Cope T/115 (TRIN, US). 
VaufIss: Rfo Guainfa, near Victor!no, river level; tree without flowers or 
fruit but with stipules; said by the local Indians to be a type of cacao with small 
smooth pods, native in the forest, 3 II 1952, R. E, D. Baker 37 (TRIN, US), 
Rfo Infrida, Santa Rosa, 300 m.; tree 15 m., trunk about 30 cm. in diameter, 
bark greenish, cortex light red, wood white, no terminal growing point, young 
shoots arising from below jorquette, cauliflorous, pod surface 10-ridged, also 
with transverse ridges, fruit pedicel about 4 cm. long with abscission layer 3 cm. 
from trunk, 25 I 1953, Bartley & Holliday T/69 (COL, TRIN, US). Left bank 
of Rio Inlrida, San Joaqufn, 200 m. from river bank, 300 m. alt.; tree 10 m., 
trunk base 15-20 cm. in diameter, one dead cymose inflorescence seen, 28 I 1953, 
Bartley & Holliday T/70 (COL, TRIN, US). Rfo Infrida, Rfo Papunana; tree 
10 m., trunk about 30 cm. in diam., bluish gray in appearance, bark red, wood 
white, three branches at each jorquette, dichotomous branch habit, inflorescences 
on upper part of trunk, 18II 1953, Bartley & Holliday T/74 (COL, TRIN, U, US). 
Amazonas: Rfo Apaporis, Jinogoj6, river level; tree 40-50 ft., 8"-9" 
diameter at base, jorquettes of 3 branches, subsequent growth from below, ultimate 
branches repeatedly bifurcating, flowers in large clusters, sepals, ligules, and 
staminodes dark crimson, 8 IX 1952, Baker & Cope 11 (COL, F, TRIN, US). 
Ibidem (boundary Amazonas-Vaup&s) between Rfo PiraparanA and Rfo Popeyaca, 
250 m., Cafio Unguyd; tree 8 m., calyx red, petals white, purplish red at apex, 
"bicco," 3-11 XI 1952, Garcia-Barriga 14380 (COL, US). 
BRAZIL: Amazonas: Sao Paulo de Olivenga, on elevated land; tree 10 m., 
12 cm. [in diam.], "cacau azul," 18 V 1945, Frdes 20942 (NY). Cidade Fonte B6a 
("introduzida pelos indios do Japuri"); tree 10-12 m., 12-15 cm. diam., "cacau 
azul," "cachu azul," 4 IV 1945, Frdes 20645 (K, USD A). B. constant, tree 10 m., 
"cacau azul," 9 V 1945, Frdes 20885 (NY). Macacacain, Rio Jutahi, Barreira 
Branca; tree 15 25 feet, small, main branches in whorls of 3, each bifid, 31 1 1875, 
Traill 62 (GH, K, P). 
6. Theobroma bernouillii Pit tier 
Figures 2, 6, 7, 15, 16, 17, 19; Map 4; Plate 5 
Theobroma bernouillii Pittier, Report. Sp. Nov. Fedde 13:319. 1914. 
Chevalier (1946) 26: 277; Le6n (1960) 314. 
Theobroma asclepiadijlorum Schery, Ann. Mo. Bot. Gard. 29:360. 1942. 
Theobroma capilliferum Cuatr. Rev. A cad. Colomb. Cienc. 6:547, figs. 8a, 
4a, pi S4. 1946. 
490 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Types.—Pittier 4105, Panama. Wedel 1535, Panama (of T. asclepia- 
diftorum). Cuatrecasas 16160, Colombia, Pacific coast (of T. 
capiUiferum). 
Tree 15-20 m. high; stem up to 25 cm. in diameter, with grayish- 
brown, rugulose bark 5 mm. thick and yellowish-white wood; sympo- 
dial growth by lateral, sub term in al, upright shoots; primary branches 
ternate, grayish brown or blackish brown, dicbotomous, the oldest 
falling off, leaving the stem naked, the terminal stem leafy, terete, 
dark brown, minutely, subappressed tomentulose, when older puber- 
ulous or glabrate; stipules linear-oblong, attenuate at apex, acute, 
pubescent, about 8 mm. long, 2.5 mm. wide. 
Leaves coriaceous, more or less rigid, entire; petiole thick, 4-12 mm, 
long, terete, densely and minutely tomentulose, transverse rimose 
when dry; blades rather asymmetrically ovate, ovate-oblong, or 
elliptic-oblong, obtusely cuneate-attenuate and mostly very asym- 
metrical at base, triplinerved, narrowed toward the apex, ending 
in a long, linear, acute tip, the margin entire or very slightly sinuate, 
slightly re volute, 13-30 cm. long, 5.5-18 cm. broad, including the 
acumen, this 1.5-6 cm. long, 2-4 mm. broad, green or pale brownish 
above when dry, shining, the main nerves filiform, prominent, the 
others reticulate, slightly prominulous, pale greenish ochraceous or 
ashy beneath, apparently glabrous but the tawny shining veins 
with very sparsely minute stellate hairs and the areolae covered 
with very appressed, white tomenturn of smaller, microscopic, 
entangled stellate hairs, the midrib and 5-7 secondary nerves on each 
side very prominent, the basal pair acutely ascending, the others 
distally curved-ascending, decurrent and anastomosing near the 
margin, the transverse tertiary nerves thin but prominent, the minor 
veins prominulous, minutely reticulate. 
Inflorescences on the main trunk usually many-flowered, borne 
on short, tuberculate, woody branches, the panicles abundant, 
4-10 cm. long, spreading, bristly, the axes slender (0.4-1.5 mm. 
thick), rigid, striolate, stellate-tomentose in the upper third or rarely 
from the base, cymoso-bifurcate, corymbiform, the branchlets rigid, 
erect at acute angle; peduncles (ultimate branchlets) capillary, 
tomentulose, 2-20 mm. long, 3-bracteolate at apex;bracteoles narrowly 
linear, 1-2 mm. long, very soon deciduous; pedicels thicker than the 
peduncles, about 5-20 mm. long at an thesis, minutely Stella te- 
tomentellous like the branchlets; buds ovoid, densely and shortly 
tomentose. 
Flowers crimson; sepals moderately thick, lanceolate-oblong, rather 
acute, shortly united at base, often first temporarily united in pairs 
but separated later, 8-10 mm. long, 3 mm. wide, reddish, with sparse 
slender, flexuous, sericeous hairs inside, ferruginous, rugulose, and 
CUATRECASAS—CACAO AND ITS ALLIES 491 
stellate-tomentose outside, minutely tomentulose at the margin 
inside, with thick, glandular hairs crowded at the insertion at base; 
petal-hood red, oblong-obovate, rounded-cucullate at apex, narrowed 
at base, 3-nerved, prominent inside, hirtellous pubescent outside, 
4-5 mm. long, about 2 mm. wide, at base 0.6 mm. wide; petal-lamina 
crimson, sessile, shortly unguiculate-articulate, moderately thick, 
rugulose, glabrous, orbicular or suboricular or elliptic, minutely 
crenulate, 2.5-4 mm. long, 3-4 mm. broad; staminal tube about 
1.5-2 mm. high; staminodes 6-9 mm. long, erect in bud, purplish red, 
sublanceolate-linear-subulate, thick, suddenly narrowed toward the 
apex, covered with minute, spreading, acute, conical trichomes; 
fertile filaments glabrous, flexuous, about 2.5 mm. long, 2-anther- 
iferous; ovary 5-sulcate-costatc, tomentose, 1.2 mm. high; styles 
five, 2 mm. long, adherent into a column but separable. 
Fruit 15-20 (12-25) cm. long, 6-7.5 (5.2-8) cm. broad, ellipsoid- 
oblong, more or less prismatic, obtusely 5-angulate, abruptly narrowed 
subacute or subobtuse at apex, umbilicate at base, more or less con- 
stricted near the base or not; pericarp thick, rigid, coriaceous at 
maturity, the epicarp and endocarp hard coriaceous, the mesocarp 
fleshy, shrinking in drying, dull brown, dense velvety-tomentose; 
seeds compressed-ovoid, 16-22 x 9-14 x 9-11 mm., the testa reddish, 
papery, the cotyledons white; pulp white, flavored, acidulous; germi- 
nation epigeous. 
This species, as here broadly considered, includes heteromorphic 
elements described as three species, two of which came from the 
Atlantic coast of Panama, and the third from the Pacific coastal 
region of Colombia. The Colombian population (T. capilliferum) is 
the best known, being represented by several collections with fruits 
and flowers, showing morphological uniformity throughout its area. 
The original T. bemouillii is known only from flowering material of 
one collection (the type), which has some minor differences from the 
Colombian plants in the shape of the leaves and details of flowers. 
The other described Panamanian species, T. ascUpiadijlorum, was 
based on discordant elements collected by von Wedel in Water Val- 
ley. Schery wrote: "Although fruiting material of this species is 
lacking, floral and vegetative characters distinguish it sufficiently to 
warrant description as a new species." The glabrous branches and 
leaves of the type specimen, which resemble those of T. cacao, do 
not belong to a Tkeobroma; they belong actually to the Lauraceae. 
The inflorescences and flowers are similar to those of T. bemouillii, 
type specimens of which were collected by Pittier not very far away. 
The question about what kind of leaves belong to the described 
flowers of T. asclepiadiflorum is answered by the collection Wedel 
681 from the same locality, Water Valley; the flowers of this collec- 
492 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
tion are identical to those of the type of T. asclepiadijlorum and the 
leaves are exactly like those of T. bernouillii. There are minor differ- 
ences between the flowers of these two types, but considering the 
vicinity of the geographic range of these two populations, it may be 
safe to consider them more forms of one species. Fruits from the 
type locality of T. bernouillii have never been collected, but Allan 
Lucas did collect fruits of T. asclepiadiflorum in Water Valley. The 
flowers of T. asclepiadiflorum are almost identical to those of T. 
capillijerumf but the fruits are smoother and not constricted. 
In view of these facts, I consider all these Colombian and Panama- 
nian plants to belong to one species; since the three forms are geo- 
graphically separated the observed differences warrant subspecific 
recognition. 
Distribution: Pacific coast and the Choc6 region of Colombia and 
the Atlantic coast of Panama. In the underforest of the tropical 
rain forest, from sea level and lowland swamps to heavily forested 
hills about 100 m. altitude. 
Key to Subspecies of Theobroma bernouillii 
1. Leaves broadly ovate or ovate-oblong, very asymmetrical, rigid, 5-12 x 2-6 
cm. Petal-lamina orbicular 2.5-3.5 mm. long; petal-hood 4-5 x 2 mm.; 
staminodes subulate, densely pilosulous, 7-8.5 mm. long; inflorescences 
long, with thin branches, the peduncles 5-20 mm. long, the pedicels about 
10 (5-20) mm. long; fruits obtusely pentagonal with smooth, conspicuous 
ridges, 12-25 x 5 8 cm., umbilicate constricted above the base. 
6c. subsp. capilliferum 
1. Leaves oblong-elliptic, slightly asymmetrical, thinner, less rigid. 
2. Staminodes lanceolate, slightly pilose, 5.5-6 mm. long, 1.5-2 mm. broad at 
base; petal-lamina suborbicular, 3 x 3-4 mm. long; petal-hood 4-5 x 2-2.5 
mm.; peduncles 2-5 mm. long; pedicels 5-8 mm. long; fruit unknown. 
6a. subsp. bernouillii 
2. Staminodes subulate, densely pilose, 9-10 mm. long, 1.5 mm. broad; 
petal-lamina elliptic, 4x3 mm.; petal-hood 5x3 mm.; peduncles 6-12 
mm. long; pedicels 10-12 mm. long; fruit oblong, 17x7 cm,, very slightly 
pentagonal, attenuate at apex, not contracted at base, with a filiform, 
impressed, furrow on each inconspicuous ridge. 
6b. subsp. asclepiadiflorum 
6a. Theobroma bernouiliii Fittier subsp. bernouillii 
Distribution.—Atlantic coast of Panama. 
PANAMA: Pro v. Coldn, in forests near Fat6, Loma de la Gloria (Nombre de 
Dios), 10-104 m., 4 VIII 1911, Pittier 4105 (US holotype; isotypes BM, BR, C, 
F, GH, K) (Photo USNH 3199). 
Further collections at the type locality are necessary to know the 
nature of the fruits. 
CUATRECASAS—CACAO AND ITS ALLIES 493 
6b. Theobroma bernouiUII Pittier subsp. asclepiadiflorum (Schery) Cuatr., 
stat. nov. 
Theobroma asclepiadiflorum Schery, Ann. Mo. Bot. Gard. 29:360. 1942; 
Le6n (1960) 316, 321, fig. (as. T. bernouillii). 
Type.—Wedel 1535, second sheet, flowers (MO, lectotype). 
Distribution.—Atlantic coast of Panama. 
PANAMA: Bocas del Toro: Water Valley, vicinity of Chiriqui Lagoon; tree 
90 feet, flowers red, 8 XI 1940, H. von Wedel 1535 (second sheet MO, lectotype); 
the first sheet of this collection (MO) belongs to the Lauraceae. Bocas del Toro, 
Water Valley, 10 IX 1940; tree 80 ft.; flowers maroon red, H. von Wedel 681 
(MO). Bocas del Toro, Water Valley, V 1949, Allan Lucas 1 (F, TURRI). 
The fruit of this subspecies is known through one specimen brought 
by Allan Lucas (TURRI), which is 17 x 7 cm., ellipsoid-oblong, very 
slightly pentagonal with 5 filiform furrows on the obtuse angles; the 
surface is slightly rugose due to the drying; the apex is shortly attenu- 
ate and the base is sub truncate and umbilicate. The shape differs 
clearly from that of the fruits of subsp. capilliferum and T. glaucum; 
it must be a mutant form geographically isolated. 
6c. Theobroma bernouillii Pittier subsp. capilliferum (Cuatr.) Cuatr., stat. 
nov. Plate £ 
Theobroma capilliferum Cuatr. Rev. Acad. Colomb. Cienc. 6:547, figs. Sa, 
4a, pi. 34. 1946; Baker, Cope & al. (1954) 13, figs. 17, 18; Le6n (1960) 
314, 317, fig. 
Type.—Cuatrecasas 16160, Colombia, Pacific coast. 
Common names.—Chocolate de monte, cacao de monte bravo, 
cacao de monte (Colombia). 
Uses.—On the Pacific coast of Colombia and in the Choc<5 area, 
the fruits are known as wild cacao (chocolate de monte, cacao de 
monte, cacao de monte bravo). Their white seeds are considered a 
high quality cacao, but the fruits remain abandoned on the trees, the 
people not making actual use of them, although monkeys and other 
animals break or pierce them, sucking the pulp or eating the seeds. 
Distribution.—Pacific Coast and the Chocd region of Colombia. 
In the under story of the rain forest, from the lowland swamps next 
to the mangroves to the forested hills about 100 m. in altitude. 
COLOMBIA: El Valle: Pacific Coast, Rio Yurumangui, Veneral, swampy 
rain forest in Quebrada del Zancudo, 5 m. elevation; tree 15 m., 25 cm. diam. at 
base, bark granulate-rugose, brown or grayish, its section 5 mm. thick, producing 
abundant mucilage; wood yellowish white; fruits 11—12 x 6 cm. (immature), 
ellipsoid-prismatic, thick, umbilicate at base, more or less constricted above the 
base, with 5 furrows or flat sides, and 5 well-marked angles, apex acute and 
slightly umbilicate, the surface minutely tomentose, greenish ferruginous, pedun- 
cles 6-10 cm. long, thick, fruits abundant, hanging on trunk, leaves coriaceous, 
rigid, green above, pale green beneath, long-tailed (Cuatr, photographs C-2202, 
2203), 10 II 1944, Cuatrecasas 16160 (VALLE, holotype; isotype, F). Pacific 
494 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Coast, Rio Cajambre, Barco, forest on hill at the right margin of Quebrada de 
Agua Clara, 40-100 m. elevation; tree 20 m., stem 25 cm. diam., leaves coriaceous, 
pale yellowish green above, cinereous beneath, distichous and pendulous, the 
lower ones larger and thicker, dry inflorescences with thin, long branchlets, 
fruiting peduncles 8-10 cm. long, 10-12 mm. thick, fruits with 5 furrows and 5 
thick angles, surface sinuate-rugose, velvety-tomentose, pale tawny, 16-20 cm. 
long, 6.5-7.5 cm. broad, umbilicate at both ends, usually constricted above the 
base, cotyledons whitish, wood yellowish white, 23 IV 1944, Cuatrecasas 17034 
(F, VALLE). Same locality and date, seedlings, Cuatrecasas 17034A (F, VALLE). 
Rfo Cajambre, San Isidro, hill forest on left margin of Quebrada de Veneno, 
about 50 m. alt.; tree 20 m., stem 20 cm. diam., bark yellowish white, leaves 
coriaceous, rigid, yellowish green above, greenish cinereous beneath, fruits hang- 
ing on trunk, 4 V 1944, Cuatrecasas 17350 (F, VALLE). Same locality,seedlings, 
Cuatrecasas 17350A (F, VALLE). Rfo Calima, Cafio de la Brea, about 5-10 
m. elev.; tree 15-20 m. high, 20-25 cm, diam. at base, crimson flowers in dense 
cushions on upper portion of trunk only, 29 VI 1943, Holliday T/142 (TRIN, 
US). Rfo Calima, Estaci6n Agroforestai, about 5-10 m. alt.; tree 12-15 m, in 
forest clearing, crimson flowers in dense cushions on upper part of trunk only, 
dry fruit up to 25 x 8 cm., ridged, 29 VI 53, Holliday T/145 (TRIN, US). 
Choc6: Llor6, young tree, sterile, 4 VIII 1953, Holliday & Bartley T/177 
(TRIN, US). Ibidem; tree C m., Holliday & Bartley T/178 (TRIN, US). Rfo 
San Juan, Remolino, young tree 2 m., sterile, 1 VIII 53, Holliday & Bartley T/ 
172 (TRIN, US). 
Nari&o: South of Tumaco, in heavy rain forest; tree 10 m., fruits on trunk, 
olivaceous, 18 X 1955, Romero Castafieda 5405 (COL). 
Map 6.—Location of known spontaneous, wild, populations of Theobroma cacao subsp. 
cacao O &nd subsp. spkaerocarpum # which may be the origin of the present cultivated 
varieties 
CUATRECASAS—CACAO AND ITS ALLIES 495 
Section 3. Theobroma 
Theobroma sect. Theobroma Figures 3, 4 
Theobroma sect. Cacao (Aubl.) Bernoulli, Uebers. Art. Theobroma 4. 1869, 
Sect. Eutheobroma subsect. Cacao (Aubl.) Pittier, Rev. Bot. Appl. 10 
(110): 779. 1930. 
Petal-laminas spatulate, long-attenuate stipitate. Petal-hoods 3- 
nerved. Staminodes linear-subulate, erect in aestivation. Filaments 
2-antheriferous. Fruit ovoid-oblong or ellipsoid, more or less pentag- 
onal, the pericarp thick, firmly fleshy glabrous. Cotyledons epigeous 
at germination. Leaves glabrous or sparsely pilose. Inflorescences 
on the trunk and on leafy branches. Sympodial growth of stem 
by orthotropic adventitious, lateral-sub terminal shoots. Primary 
branches in 5's, persisting in age. 
Type species.—Theobroma cacao L. 
7. Theobroma cacao L. 
Figures 1, 2, 3, 5, 6, 20, 21, 22, 23, 24, 25, 26; MAP 6; PLATE 6 
Theobroma cacao L. Sp. PI. 2:782. 1753; Syst. Nat. ed. 12, 2:508. 1767; 
H. B. K. (1823) 316; Richard (1845) 183 (1845a) 73; Bernoulli (1869) 5, 
pi If Baillon (1884) 792-795, figs.; Schumann in Mart. (1886) 72, pi 7; 
Jumelle (1899) 11, figs. 1-9; Preuss (1901), pis. 1, 2; De Wildeman (1902) 
91, figs. 16, 18, #0, SI. 1902; Standley (1923) 805; Ducke (1925) 130; 
(1940) 268, pi. 1, fig. 1, (1954) 11; Fawcett & Rendle (1926) 158-160, 
fig. 60; Uittien in Pulle, Fl. Surinam 3:45. 1932; Ciferri (1933) 604; 
Standley (1937) 688; Chevalier (1946) 269-274, pi. 5; Standley & Stey. 
(1949) 423; Hoidridge (1950) 4; Addison & Tavares (1951) 25, pi. 7, pi. IS, 
fig. 7; Lem6e (1952) 379; Baker, Cope & al. (1954) 9-11; Cuatrecasas 
(1956) 653; Le6n (1960) 311-313. 
Cacao Clusius, Exot. Libr. Dec. 55. 1605; Ray (1688) 1670; Sloane (1696) 
134; Tournef. (1700) 660, pi 444; 1700; Merian (1705), pi 26; 
Sloane (1725) 15, pi. 160; Ray (1733) 158; Weinm. (1739) 1-11, pi 277, 
278; Geoffr. (1747) 409; Catesb. (1747) 6, pi 6; Blackwell (1739) 373; 
HernAndez (1630) 79, (1946) 908-914. 
Amygdalis similis guatimalensis, Avellana mexicana Bauh. Pinax Theat. 
Bot. 442. 1623. 
Arvor cacavifera americana Pluk. Almagest. Bot. 40, pi 268. 1696. 
Theobroma foliis integerrimis Linn. Hortus Cliff. 379. 1737. 
Cacao guianensis Aubl., PI. Guian. 2:683, pi 275, figs. 1-15. 1775, pro parte 
(tantum flores). 
Cacao saliva Aubl., PI. Guian. 2:689. 1775; Lam. Encycl. Meth. 1:533, pi. 
635. 1797. 
Cacao minus Gaertn. Fruct. & Sem. 190, pi. 122. 1791. | 
Cacao Theobroma de Tussac. Fl. Antill. 1:101, pi IS. 180&. 
Theobroma iniegerrima Stokes, Bot. Med. 4:83. 1812. 
Theobroma caribaea Sweet. Hort. Britt. 67. 1830 (nom. nud.) 
Theobroma peniagona Bernoulli, Uebers. Art. Theobroma 6-7, pi 2. 1869; 
Preuss (1901) 199, 255, pi. S, 5; De Wildeman (1902) 94; Standley & Stey. 
(1949) 427. 
Tkeobroma leiocarpa Bernoulli, op cit. 6, pi 2; Standley (1937) 688; Standley 
& Stey. (1949) 426. 
496 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Theobroma Saltzmanniana Bernoulli, op. cit. 7, pi. 2, 
Theobroma Kalagua De Wild., Bull. Herb. Boiss. 7:957, pi. 1 i. 1899 (tantum 
folia, sed lectotypus), 
Theobroma sativa (Aubl.) Lign. et Le Bey, Bull. Soc. Linn. Norm. V, 8:263. 
1904; Chevalier (1946) 270. 
Theobroma sphaerocarpa Chevalier, Veget. Util. Afr. Trop. Fr. 4:12. 1908. 
Theobroma sapidum Pittier, Bol. Soc. Venez. Cienc. Nat. 1:183. 1932, nom. 
nud. 
Theobroma cacao var. typica Ciferri, Monogr. 604. 1933. 
Theobroma cacao var. Uiocarpa (Bernoulli) Ciferri, Monogr. 604. 1933. 
Theobroma cacao var. typica x T. cacao var. leiocarpa, Ciferri, 604. 1933. 
Theobroma cacao forma leiocarpum (Bernoulli) Ducke, Rodriguesia 4:274. 
1940. 
Theobroma sativa var. leucosperma Chevalier, Bull. Inter. Bot. Appliq. 26: 
270. 1946. 
Theobroma sativa var. melanosperma Chevalier, loc. cit. 
Theobroma cacao subsp. leiocarpum (Bernoulli) Cuatr. in Macbr. Fl. Peru, 
Field Mus. Publ. Bot. 13 (3A): 654. 1956. 
Theobroma cacao subsp. sativa (as (Lam.) Lign. & Le Bey) Le6n in Hardy's 
Cacao Man. 312. 1960. 
Theobroma cacao subsp. pentagona (Bernoulli) Le6n, loc. cit. 
Type.—Sloano Herbarium vol. 5, p. 59 (BM, lectotype); Paratype: 
Tourneforfc pi. fruit-lectotypc, 
Tree usually 4-8 m. high, rarely taller (up to 20m.), with the 
growth of the sympodial stem by subterminal, lateral, upright shoots 
(chupons); primary branching by successive whorls of normally 
spreading branches; young branchlets terete, grayish green or brown- 
ish, densely or sparsely pubescent, with slender, patulous, acute, 
simple or furcate hairs 0.1-0.3 mm. long, later glabrate, more or less 
striate, rugulose and sparsely lenticellate; stipules subulate, very 
acute, 5-14 mm. long, 0.5-1.5 mm. wide at base, pubescent or pu- 
berulous, deciduous. 
Leaves coriaceous or chartaceous, more or less rigid, alternate, 
distichous on the normal branches, green; petioles pubescent or 
tomentose, with simple, acute, slender, rather dense, spreading hairs, 
thickened pulvinate at the ends, 1.5-2 (1-3) cm. long, on orthotropus 
stems 3-10 cm. long; blades subobovate-oblong or elliptic-oblong, 
slightly asymmetrical, rounded or obtuse at base, attenuate and 
cuspidate at apex, acute or subacute, usually entire or slightly and 
irregularly sinuate, green above, pale when dry, glabrous except for 
the pubescent or puberulous midrib, the midrib linear, prominent, 
the secondary nerves filiform, the fine veins reticulate, often slightly 
prominent, lighter green beneath, glabrous or with very sparse, 
minute, simple, furcate or stellate hairs, rarely puberulous, the 
midrib thick and prominent, the secondary nerves 9-12 each side, 
prominent, subpatulous, then ascending, near the margin curving, 
slendering, anastomosing, the tertiary nerves prominent, the minor 
veins reticulate and prominulous, 15-50 cm. long, 4-15 cm. broad, 
the acumen 1-2.5 cm. long. 
CCATRECASAS—CACAO AND ITS ALLIES 497 
Figure 20.—a-i, Theobroma cacao subsp. cacao fma. pentagonum (Cuatr. 26004): A, B, c, 
petal from inside, outside and laterally, X 5; D, androeeium, X 5; E, stamen, X 10; 
F, anther, X 20; g, pistil, X 10; h, bud, X 2; i, sepal from inside and outside, X 2. 
k-q, T. cacao subsp. sphatrocarpum (Cuatr. 7756): k, l, m, petal from inside, outside 
and laterally, X 5; N, androeeium, X 5; o, stamen, X 10 and anther, X 20; p, pistil, 
X 5; q, sepal from inside and outside, X 2. r-y, T. cacao subsp. cacao (Nelson 2490): 
&, 8, T, petal from inside, outside and laterally, X 5; androeeium, X S;v, stamen, X 10 
and anther, X 20; w, pistil, X 5; bud and pedicel, X 2; v, sepal from inside and outside, 
X 2. 
498 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Inflorescences on the trunk and branches, usually borne on small 
tubercles, the cymose branchlets short, knotty, persistent, the cymose 
peduncles 1-3 mm. long, stellate-pubescent, hirtellous and with, 
scattered glandular trichomes; bracts ovate or ovate-oblong, amplec- 
tant, pubescent; bracteoles ovate-oblong, acute or subacute, 0.5-1.2 
mm. long, pubescent, deciduous; pedicels capillary, rigid, pale green, 
whitish or reddish, 5-15 mm. long, hirtellous with rather dense, thin, 
Figure 21.—Sketches of fruits of Tkeobroma cacao from classical and recent literature: 
a, in Tournefort plate 444, clearly representing a "criollo type"; b, in Sloane, pi. 160; 
c, in Chevalier 1946, pi. 5 (7*. sphaerocarpum Chev.). 
CUATRECASAS—CACAO AND ITS ALLIES 499 
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Figure 22,—a, Fruit of Theobroma cacao, cultlvar "cundiamor" (Cuatr. 26492 from 
Colombia)* b, Cross section of same showing structure of pericarp and arrangement 
of seeds each surrounded by their pulp, c, T. cacao subs p. cacao, cultivar (tcriolloM 
("Caldas") (Cuatr, 26006 from Colombia), d, Section of same. All X J4* 
500 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
acute, patulous, stellate or furcate hairs and sparse pluricellul&r, 
glandular, capitate trichomes; buds white, whitish green, lilac, or 
reddish, ovoid or oblong-ovoid, acute, 5-7 mm. long, subglabrous or 
sparsely puberulous with slender stellate hairs and thicker, glandular, 
Figure 23.—a, Fruit of Theobroma cacao subsp, cacao fma. pentagonum (Cuatr. 26004 from 
Colombia) with some degree of introgression from fma. cacao, b, transection of the 
same showing the great reduction of the structure of pericarp (mesocarp represented 
by isolated bundles), c, T. cacao subsp. cacao fma, pentagonum, typical (Cuatr. 
26540 from Costa Rica), d, transection of the same, showing the pericarp reduced to 
one layer. All X CuItivar'Magarto." 
CUATRECASAS—CACAO AND ITS ALLIES 501 
Figure 24.—a, Fruit of Theobroma cacao subsp. spkaerocarpum, cultivar "amelonado" 
(Cuatr. 25805 from Venezuela, Barlovento). b, transection of the same, c, fruit of 
T, cacao cultivar "angolcta" (Cuatr. 26494 from Colombia), d, transection of same. 
A11X 
stipitate trichomes; sepals thick-membranaceous, lanceolate or oblong- 
lanceolate, acute, white, greenish, white, pale violaceous or reddish, 
slightly 3-nerved, shortly (0.5-1 mm.) united at base, 5-8 mm. long, 
1.5-2 mm. wide, minutely tomentose at margin, outside sparsely 
pubescent with stellate and glandulose hairs, or glabrous, inside 
glabrous or with rare glandular trichomes; petals contorted in aestiva- 
tion, thick-membranaceous, the hood 3-4 mm. long, 5-2 mm. wide, 
502 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
* .<:iM 
* ' \J- " ft 
- •:£?- 
Figure 25.—a, Fruit of Tfuobroma angustifolium (Cuatr. 25790). a, fruit of T. cacao 
subs p. cacao fma. lacandonensi (Miranda 9299), c, 7*. cacao subs p. sphacrocarpwn 
cultivar "calabacillo" (Cuatr. 25805P from Venezuela), d, section of the same, e, 
fruit of T. cacao subsp. cacao f. Uiocarpum from the original drawing by Bernoulli, f, 
T. hylatum (Araque & Barkley 18C745). All X %• 
CUATRECASAS—CACAO AND ITS ALLIES 503 
Figure 26.—a-c, Showing the characteristic elongated, bipulvinate petiole in Tkeobroma 
cacao: a, T. cacao from orthotropic stem (Steyermark 54143); b, T. cacao from lateral 
branches (Cuatr. 7756); c, 7". cacao from young, lateral branches (Steyermark 49218). 
d, exceptional, broad, rounded leaf of T, cacao, a cultural mutation (Leon 5078). E, r, 
7*. microcarpum (Froes 23963 and Holliday & Cope 125). c, young shoot showing 
pubescens and stipules of T. microcarpum (Holliday & Cope 125). H, T. gileri (Giler 
162). o,X 3; leaves, X H* 
680-6GB—64 9 
504 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
obovate, rounded at apex, white, 3-nerved, the nerves papillose inside, 
the lateral ones very thick, usually purplish or red, the medial promi- 
nent only upwardly; petal-lamina pale yellowish, 1.5-2.5 mm. long, 
1.5-2 mm. wide, obovate or rhombic-obovato, attenuate at apex, 
acuminate or subtruncate and shortly mucronate, rarely blunt, entire 
or sinuate at margin, pedicellate at base, the pedicel linear, 3-nerved 
and sometimes 4- or 5-nerved at apex, 2-4 mm. long, 0.2-0.5 mm. 
wide; androecium tube rather thick, 1.2-1.5 mm. long, glabrous; 
staminodes 4-6 mm. long, 0.6-0.7 mm. broad at base, narrowly 
subulate, very acute, erect in bud, erect or somewhat flexuous in 
an thesis, the middle vein thick, angular, red or purplish, minutely 
papillose-pilose, the thin margin whitish, more or less revolute, ciliate, 
with slender, flexuose simple hairs; stamens diantheriferous, the 
filaments glabrous, patulous or recurved, 1.5-2 mm. long, the anthers 
about 0.4 mm,, with rounded cells; ovary oblong-ovoid, obtusely 
pentagonal or 5-ridged, about 1.5 mm. high, 0.8 mm. thick, glabrous 
or usually glandular, covered with more or less copious white or 
reddish, pluricellular, stipitate glands; styles 5, glabrous, adherent, 
1.5-2.5 mm. long. 
Fruit subbaccate, variable in shape, from globose to fusiform and 
acute, and with a very smooth to a strongly ridged and rugose or 
verrucose surface; pericarp consistently fleshy and thick (5-15 mm.), 
and usually made of two, more or less conspicuously different, 
carnose layers (epicarp and endocarp) separated by a thin, 
ligneous membrane (mesocarp), the endocarp limited by a firm 
epidermis inside, the inner wall of the shell, the epicarp sometimes 
differing in color and firmness, rich in mucilaginous cells, limited 
outside by the hard epidermis of the fruit, the mesocarpic membrane 
sometimes reduced to isolated bundles of fibers or lacking, the endocarp 
also sometimes lacking. 
Seeds (20^0) usually arranged in 5 rows, but sometimes when 
large arranged in 4 or 3 rows, the five radial walls initially separating 
the 5 cavites in the earlier stages reabsorbed long before the maturation 
of the fruit; seeds ovoid, ellipsoid, amygdaloid, more or less complanate 
through mutual pressure or almost round in cross section, variable in 
size (20-40 mm. long, 12-20 mm. broad), the integuments brown, 
subcoriaceous, the surrounding pulp white, sweet, the cotyledons 
white, purplish, violet, or intermediate in color, 
Theobroma cacao is variable in its characters, especially with regard 
to the color, size, and shape of the parts of the flower and the fruits. 
These are variations to be expected of an ancient crop spread 
throughout a very wide area. 
Based on some of these more or less consistent variations, Bernoulli 
described three species, T. pentagonum, T. leiocarpum, and T. saltz- 
CUATRECASAS—CACAO AND ITS ALLIES 505 
mannianum, and Chevalier T. sphaerocarpum. The shape of the fruit 
is the main defining character, except for T. salt zmannianum, which 
was based on petal characters, probably from an abnormal specimen. 
The few floral characters given for the other three species are irrelevant 
or inconstant. Tkeobroma pentagonum is defined by having elongate, 
gradually and acutely narrowed, warty pods which are strongly pen- 
tagonal and 5-ridged; it has white seeds. It was described from the 
Atlantic coast of Guatemala and is called "cacao lagarto." Tkeobroma 
leiocarpum was characterized by ovoid pods, almost smooth, with five 
very shallow furrows and a glabrous ovary; the color of the seeds 
was not stated; it was found in plantations on the Atlantic coast of 
Guatemala, and stated to be rather rare. Tkeobroma sphaerocarpum 
was described from cultivated specimens on Sao Tom6 island, 
western Africa, characterized by its nearly spherical, slightly 10- 
furrowed, almost smooth or slightly rough pods, and violet cotyledons. 
Schumann considered the three Bernoulli species as mere local varia- 
tions of T. cacao and therefore unworthy of taxonomic consideration. 
Some authors have followed Schumann in this view, but there is a 
tendency to accept T. pentagonum as a different species, because of its 
characteristic fruit form and thinner pericarp. 
For many years there has evidently been confusion in the taxonomy 
of cultivated cacaos, the main problem being insufficient knowledge 
of the wild populations of T. cacao. There are many citations of 
places in Central and South America where T. cacao is said to have 
been found wild, which may have been true in some cases, but not in 
others where we may be dealing with the remains of abandoned 
plantations. The discovery of wild cacao by Stahel in the rain 
forests of Mamaboen Creek in Surinam (confirmed later by Myers) 
and a few other places very distant from populations, is very significant; 
the cacao is of the Amelonado type. Ogilvie also found it in abundance 
on Black Creek, a branch of the Essequebo River, and along the 
Berbice River in British Guiana. He found it along the Rupunumi 
River at Rewa and Quitaro Creeks, at Kuduwini Kassikedju or 
Dewar Wow, etc., on the upper Essequebo River, and also on the 
Quitari River (according to Myers). Robert Schomburgk also cited 
wild cacao in several places in British Guiana on the Cutari, at the 
head of the Correntyne River, and also at Quitaro, and Richard 
Schomburgk found it on the Upper Pomeroon River. Wild cacaos 
also have been found, according to Stahel, in the Upper Oyapok in 
French Guiana (Myers), all of the Amelonado type, implying that 
this population of the Guiana highland area westwards to the Amazon 
valley may have been the home of this cultivated variety. In Brazil 
there has been found wild cacao near the Guianas and, according to 
Ducke, at the upper Rio Branco, northeast of Obidos, and at Francez 
506 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
on the middle Tapaj<5z; he adds that the area of spontaneous T. cacao 
includes a greater part of the Hylaea and that it is of the form "leio- 
carpum" Preuss (1901, p. 247) found wild cacao in eastern Ecuador, 
also of the Amelonado type. Huber found spontaneous cacao in the 
upper Purus river, and along the Rio Ucayali, Peru. Pound also 
claimed to have found spontaneous cacaos in the upper Amazon 
basin. The spontaneous cacao trees found by the Anglo-Colombian 
Expedition in the forests of Cagu&n (Caqueta) and Rio San Miguel 
offer some doubt about their indigenousness. I found myself, in 
1939, in the rain forests of Rio Guaviare wild trees of T. cacao, tall 
and well developed, but we have to be cautious in accepting them as 
indigenous, because the Guaviare River is said to have had more 
extensive plantations on its banks in earlier times. An interesting 
find was made in British Honduras by Sampson, who encountered 
wild trees of the Criollo type in the Southern forests. In Mexico, Miranda 
recently found a completely indigenous cacao tree in a forested region 
of Chiapas (Selva Lacandona), very distant from any population; the 
natives (Lacandones Indians) reported that other trees of such cacaos 
are found scattered; this cacao has a somewhat scandent stem and 
produces small, elongated, pointed, slightly 10-ridged and rugose 
fruits. Standley and Steyermark (Fl. Guat. 426) say that in the 
lowlands of Guatemala sometimes cacao is found more or less wild in 
the forest, especially in Alta Verapaz, and that it is not improbable 
that it is native in the wet North Coast region. 
Many forms of the cultivated cacaos have received local or regional 
names which after the many introductions of cultivars from one 
country to another have brought confusion to the complicated system 
of cacao types. Morris was the first to make a classification of 
cultivars, arranging them in two main groups: Criollo and Forastero. 
Hart (see page 396) modified this, making three groups (Criollo, 
Forastero, and Calabacillo), later separating T. pentagonum (cultivar 
"Lagarto") into another group, as a different species from T, cacao. 
The classification of Hart has been followed by many authors of 
cacao treatises until recent times when van Hall, simplifying, went 
back to the basic two groups of Morris: Criollo and Forastero. 
Pittier, a botanist with much experience both in Central and South 
America, recognized the existence of two different, original, spon- 
taneous forms of cacao, Criollo with elongated, ridged, pointed fruits 
and white cotyledons, and Forastero, with short, roundish, almost 
smooth fruit and purplish cotyledons; he believed that they corre- 
sponded with two different species, T. leiocarpum generally spread 
throughout tropical South America, where it is still found spontaneous, 
and T. cacao, spontaneous in Central America and which was the 
h 
CUATRECASAS—CACAO AND ITS ALLIES 507 
source of the prehistoric cultivation and selection of cacao, all of 
the Criollo type, in Mexico and Central America. Introduction of 
Criollo types in South America and West Indies and conversely of 
the smooth type into West Indies and Central America created the 
cross-varieties which multiplied with the years. Although there is 
much speculation in this, the theory is a workable and reasonable 
one, because the available historical data favors the recognition of 
the earlier Central American and Mexican cacaos as being of the 
Criollo type. The finding of spontaneous cacao of this type in Chiapas 
and British Honduras also supports this theory. Another favorable 
fact is the uniformity of the Venezuelan Criollo, supposed to have 
been introduced from Central America to Venezuela in the earlier 
times of the Spanish conquest. 
Soria, after visiting (1961) important plantations in Mexico where 
new Forastero types have been abundantly introduced in recent times, 
recognized that in Mexico before 1900 varieties of the Criollo type 
almost exclusively were cultivated. He observed in large, old 
(more than 50 years) plantations a great variation in the Criollo type, 
but the seeds were always white and the pods, variable in shape, 
were always pointed, with surfaces varying from tuberculate to rugose, 
from light green, through green, to reddish; the trees were small, 
slow growing, and often with fewer branches (5-3 in each whorl) than 
is normal in the species; the petal-laminas are bright yellow. 
Pittier's theory was very much welcomed by botanists and cocoa 
experts; Chevalier supported it, and Ducke also in its basic idea. 
Cheesman adhered to it at first (1932), but later (1944) developed a 
new theory that all cultivated Criollo cacaos came "from an offshoot 
of a general cacao stock in the headwaters of the Amazon carried over 
the Andes into southern Colombia, and developed many of their 
present characters in association with man." But historical knowledge 
at present can only closely relate cacao to Central American man, 
especially to the Mayans and not to the South American Indians. 
Central American Indians undoubtedly developed the art of planting 
and selecting of cacao through several thousands of years, finally 
obtaining the high quality produce which the Spaniards found at the 
time of the conquest. 
It may be assumed that in early times a natural population of 
Theobroma cacao was spread throughout the central part of Amazonia- 
Guiana, westward and northward to the south of Mexico; that these 
populations developed into two different forms geographically sepa- 
rated by the Panama isthmus; and that these two original forms, when 
isolated, had sufficiently consistent characters to be recognized as 
subspecies. As they intermingle readily by crossing, giving fertile 
508 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
and robust hybrids, they cannot be considered distinct species. Both 
types in cultivation have originated many mutations, some of them 
persistent, thanks to ecological adjustment, selection, and isolation. 
Because the cultivation and selection has been very active for some 
thousand years in the Central American-Mexican area, it is in these 
areas where we find the richest variety of forms. When the sub- 
species (with their different forms) interbred the products gave the 
great and confusing variety of existing types. 
Another theory is that most of the stable forms of cacao might 
have originated by mutations from a widespread, uniform original 
specific type. In such a way the forms pentagonum, leiocarpum, 
Criollo, Cundiamor, Angoleta, Nacional Ecuador, Trinitario, etc., 
and their variants could have originated. This theory does not ex- 
clude hybridization as a factor in the multiplicity of forms, but its 
influence would be secondary instead of basic, as postulated by the 
Pit tier theory. 
Types of Theobroma cacao.—The only specimen of T. cacao in 
the Linnaean herbarium is the number 934-1, but this specimen is 
posterior to the publication of Species Plantarum (1753), since it bears 
the annotation "PI. Surin. 1775. p. 13." The type has to be sought 
among the bibliographic references of Linnaeus. In Species Plan- 
tarum, Linnaeus refers to Hortus Cliffortianus. At my request, 
Mr. Sandwith was kind enough to examine the Hortus Cliffortianus 
herbarium at the British Museum and found no specimens of Theo- 
broma in it. However, Hortus Cliffortianus page 379 seems to give a 
key to the matter, where Linnaeus writes "Flores a nullo bene depicti, 
multo minus descripti sunt, . . . Sloane mihi inspiciendi copiam 
fecit, . . and gives the quotation "Hist. Jam. pi. 160 " Linnaeus 
was especially preoccupied with the flowers; he wanted to know 
exactly the structure in order to be able to place Theobroma in the 
right place in his sexual system; the previous literature did not give 
him the answer, and neither did the Sloane plate. However, from the 
above paragraph we may infer that Sloane sent him flowers at his 
request, surely very few, which may be the reason why there are none 
of them in the Linnaean herbarium; from Sloane's flowers Linnaeus 
found the floral structure of Theobroma by himself and drafted his 
diagnosis. The original flowers (surely dissected) having disappeared, 
the corresponding specimens in the Sloane Herbarium have to be 
considered the isotype. Messrs. Dandy and Sandwith found the 
specimens in the Sloane Herbarium at the British Museum. Mr. 
Sandwith writes: "We found the corresponding specimen in the 
Sloane Herbarium and it is obviously the source of (in fact part of) the 
plate, though not identical; there are leaves, detached flowers, frag- 
CUATRECASAS—CACAO AND ITS ALLIES 509 
ments of fruit and one seed; it is the vol. 5 p. 59." There is the possi- 
bility that Sloane lent Linnaeus these specimens, but even if this were 
not the case, we may assume that Linnaeus used flowers taken from 
this specimen which would thus be an isotype; therefore, the sheet 
page 59, volume 5, of the Sloane Herbarium is to be chosen as lecto- 
type for the flowers and leaves. According to Sand with and Dandy 
"there is also what appears to be a duplicate, leaves only, in the 
Flukenet Herbarium volume of Herb. Sloane, The leaves of both 
specimens are reddish brown and glabrous beneath with reticulate 
tertiary veins." 
Sloane's plate 160 is a para type, but another even more important 
paratype is Tournefort's plate 444. Tournefort is the only reference 
given by Linnaeus in his original description of the genus in Genera 
Plantarum (1754), and its citation precedes the reference to Sloane 
in Hortus Cliffortianus. I propose it as the lectotype for the fruit, 
because the Tournefort drawing is perfectly defined, leaving no doubt 
that the characters are of the Criollo type. On the other hand, the 
fruits shown on Sloane's plate 160 are not unquestionable, even 
though they are very pointed; they are among the variations found 
in Criollo populations. But the reasons which compel me to consider 
the flower specimens in the Sloane Herbarium as the lectotype do not 
apply to the fruit, the origin of which may have been different from 
the origin of the flowers and leaves, for Sloane collected in several 
places and countries. A fruit typification by the Tournefort plate 
fits well the definition of T. cacao L. sensu stricto given by Pittier 
and later authors. It is obvious that all these authors described 
cultivars and that the cacao described by the earlier authors was of 
the type Criollo, as can be inferred from the illustrations, and also 
from the descriptions of Hernandez, Pison, Plukenet, Merian, 
Weinmann, Tournefort, Catesby, and Gaertner. 
Synonyms.—T. pentagonum Bernoulli was characterized by the 
shape of the fruit and by smaller flowers. Last character is inconstant 
but the fruit form is a very particular one and constant, fruit always 
easily recognized. This type was described in vivo by Bernoulli 
from the Atlantic coast of Guatemala; there is no type specimen 
of the fruit, but it was well drawn (PL %,jig. Ill) and the drawing may 
be considered the type. It is my belief that T. pentagonum is just a 
cultivar. It crosses easily with other forms of cacao giving inter- 
mediate products. Gross morphological study also supports this 
view. The pericarp in Tkeobroma is composed of three visible layers, 
one of these being more or less consistently woody; in T. cacao the 
woody layer is the middle one, the mesocarp. It seems that in 
cultivation there is a tendency for the pods to change, the shells 
510 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
becoming thinner in the beat quality Criollos. This reduction is 
extreme in T. pentagonum which lacks the firm mesocarp and the 
fleshy endocarp, the whole pericarp being only half as thick as in 
other cultivars; pentagonum trees are also weaker than others. I 
have no doubt that pentagonum is a fixed product of mutation selected 
for better fruits. I must agree with Soria (1959) when he writes 
"Pentagona is nothing more than one of the extremes in the variability 
of the complex of types forming the species T. cacao" The name 
must be kept but only at the rank of forma. 
Theobroma leiocarpum was also described by Bernoulli from living 
specimens in cultivation on the Atlantic coast of Guatemala; there are 
no type specimens of the fruits recorded, and so the published drawing 
(PL fig. II) must be chosen as the type. This plant was charac- 
terized by a glabrous ovary and smaller flowers (an insignificant 
feature) and by ovoid, shallowly 5-sulcate, almost smooth fruits. The 
color of the cotyledons was not mentioned. For a long time this form 
was identified with the widespread South American "Calabacillo" or 
"Amelonado" fruit types, especially since Pittier published his theory, 
it being supposed that the Venezuelan "Calabacillo," with thick- 
shelled, rounded or ellipsoid, obtuse, slightly 10-furrowed fruits, and 
violet cotyledons, was T. leiocarpum. All workers followed this nomen- 
clature, myself included. It was Cheesman (1944, p. 14) who called 
attention to the differences between Calabacillo and Central American 
T. leiocarpum. The Bernoulli cacao has thin, ovoid attenuate shells, 
with only 5 furrows, and plump seeds which are probably white or 
light violet. Preuss had written previously that the seeds were 
different. I saw Calabacillo in Nicaragua with very light-violaceous 
seeds. The recent observation by Soria (1961) in Mexico of the great 
variety in external form of Criollo in an old plantation of Criollo 
makes it clear that T, leiocarpum is a mere segregate form or mutant 
from the Criollo type of T. cacao, originating in a similar way to T. 
pentagonum. I consider it only as a cultivar. 
Theobroma sphaerocarpum was described by Chevalier from cultiva- 
tion in Isla Sao Thom6, in the South Atlantic Ocean west of tropical 
Africa, conforms perfectly with the "Calabacillo" cultivar of Venezuela 
and other South American plantations. It is an extreme form of the 
widespread South American subspecies. The type is a preserved fruit 
in the Museum in Paris. This name has to take the place of the sub- 
species which Pittier and later authors have called T. leiocarpum. 
Cacao saliva Aubl. is a nomenclatural synonym of T. cacao, and 
cannot be used as a substitute for the latter. Any imprecision implied 
by the binomial T. cacao is implied also in Cacao saliva, and Theobroma 
saiivum. 
CUATRECASAS—CACAO AND ITS ALLIES 511 
Theobroma sapidum Pittier may well have been unintentionally 
published as a lapsus calami for T. sativum; it corresponds to T. 
cacao sensu stricto of Pittier, restricted to the Criollo type. But the 
binomial is a nomen nudum, because it was not formally published, 
not being accompanied by a description and with no indication of any 
type. 
Cacao minus Gaertn. was published without mention of specimens 
or locality. It agrees well with some forms of the Criollo type. It 
cannot be T. pentagonum because in T. pentagonum the ridges are 
always very prominent and smooth. The type of the binomial 
consists of two fruits identical with the original drawing, labeled 
Cacao minus, Gaertner at Paris. 
Cacao theobroma de Tussac, T. integerrima Stokes, and T. caribaea 
Sweet are nomenclature! synonyms. 
Theobroma saltzmannianum was established by Bernoulli, the emar- 
ginate petal-laminae being the only difference from T. cacao (ligulae 
lamina anguste obovata, apice truncata emarginata). According to 
Bernoulli the shape of the ligula was a constant character in T, cacao 
and T, leiocarpum; having found at Kew some specimens collected by 
Salzmann near Bahia, in which he saw the petal-lamina emarginate, 
he did not hesitate to make a new species. Schumann could not 
identify this character in any of many flowers collected in Bahia by 
Salzmann, and inferred that Bernoulli had examined some exceptional, 
teratological flower. Kombouts in 1948 (Kew Bull. 1948: 104) 
studied in detail the type specimens at Kew and arrived at the same 
conclusion as Schumann, that Bernoulli did base his species on an 
accidental character. Chevalier had already expressed the same 
view (1946, p. 270). I also can confirm the above opinions after 
having examined several Salzmann collections at different times and 
the type specimen in 1954 at Kew. 
Theobroma sagittata Pavon was published by Chevalier in his revision 
(1946: 274) as a microspecies of the complex of T. cacao. The bino- 
mial, however, is not validly published for lack of the required Latin 
diagnosis. Moreover, I have identified the type specimen, which is 
preserved in Paris, as a 3-leaflet fragment of a leaf of Herrania nitida 
(Poepp.) Schultes. Theobroma hastata, a name mentioned by Cheva- 
lier in the footnote on page 273, presumably is a lapsus calami for 
T. sagittate,. 
The varieties leucosperma and melanosperma published by Chevalier 
without reference to any type specimens are fortunately not validly 
published for lack of Latin diagnoses. It would be very difficult to 
ascribe these two supposed varieties to any recognized taxonomic 
entity relying only on the seed color. 
512 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Common names.—Cacao, cocoa, cacao dulce, cacao criollo, cacao 
calabacillo, cacao forastero, cacao amelonado, cacao trinitario, cacao 
lagarto, and many other adjectives according cultivar varieties and 
regional or local cultivar forms. Also: Bizoya, yagabizoya (Reko), 
deghy (Otomi), caocauatzaua (Zoque), kako (Mixe), cahequa (Tar as- 
cdn), Chudechu (Otomi) in Mexico after Standley; cacahuatl, caca- 
hoatl, cacahoaquahuitl, quauhcacahoatl, mecacahoatl, xochicacahoatl, 
tlacacahoatl, cacahoacuahuitl, cacaotlquahuitl (Nauhatl), kicou, kicob, 
cuculat, pacxoc, cucuh, caco in Costa Rica and Guatemala; cacao 
chuncho in Peru. For Costa Rica and ChiriquI, Pit tier (1902) gives 
the following Indian names: ko (Terreba), ko6 (Brunka), kajo (Gua~ 
tuso), ku& (Quaimi), kno (PenonomS), dol6 (Doraske), tsiru (Bribri), 
(Cabicara); according to him the Bribri Indians use the following 
names for some varieties of cacao: muru-uak, tsipa-uak, xi-uak, and 
betstin-uak; Standley (1937) mentions, also for Costa Rica: kuk 
(Rama), tsiru-kuru (Cabecara), kao-kra (Brunka) and kau (TiHbi). 
Cacao silvestre, cacao de monte, wild cacao, are names often used in 
different countries whenever a cacao tree is found apparently spon- 
taneously growing. It is a fact that although the other species have 
native Indian names, for Tkeobroma cacao only the name "cacao" is 
recorded from the whole of South America, whereas this species has 
many indigenous names in Central America. 
Subspecific divisions.—A correct classification of all forms will 
only be possible after careful and extensive genetic research. In the 
meanwhile we cannot do more than use a provisional, conservative 
approach, confining the nomenclature to names already used. 
The summarized classification given below of cultivated varieties 
follows Morris, who was the first to give status to the most popular 
common names of cocoa cultivars; the adaptation in general use made 
by van Hall is followed in this paper, with few modifications. See also 
my reviews of Preuss (1901), and Hart (1892, 1900 and 1911) in the 
Historical Sketch above. The reader will find more extensive infor- 
mation on this subject in special treatises (van Hall, Baker in Urqu- 
hart, Hart, etc.). 
Key to subspecies and forms of Theobroma cacao L. 
1, Fruit elongate, claviform, fusiform or ovoid-oblong, tapering and pointed, more 
or less strongly 10-costate or 5-costate and verrucose; pericarp moderately 
thick, the woody mesocarp thin; seeds ovoid or ellipsoid, usually rounded 
in cross section; cotyledons white or yellowish white . . 7a. subsp. cacao 
2. Fruit 10-costate .... 1. f. cacao, lacandonense and unnamed forms. 
2. Fruit claviform, strongly 5-costate, the ridges prominent and smooth, the 
sides strongly verrucose, the pericarp thinner, lacking the woody mesocarp 
and endocarp 2. f. pentagonum 
2. Fruit ovoid, shallowly 5~furrowed, almost smooth, obtusely attenuate at 
apex ............. 3 f. leiocarpum 
CUATRECASAS—CACAO AND ITS ALLIES 513 
1. Fruit ellipsoid, almost globose or more or less oblong, rounded at both ends, 
smooth or slightly verrucose, more or less shallowly 10-fur rowed; pericarp 
very thick, the woody mesocarp firm; seeds ovoid, more or less compressed; 
cotyledons purplish or dark violet 7b. eubsp. sphaerocarpum 
7a(l). Theobroma cacao subsp. cacao Fiqubes 20, 21, 22 
Cacao minus Gaertn. 1791. 
Theobroma sapidum Pittier, 1932. 
Theobroma cacao var. typica Ciferri, 1933. 
Theobroma saliva var. leucosperma Chevalier, 1946. 
Type.—Sloan e Herb, (flowers, leaves) (BM, lectotype); Tournefort 
pi. 444 (fruit-lectotype). 
Common names.—The leading name is cacao criollo or criollo. 
DISTRIBUTION,—Originally from Mexico and Central America, and 
cultivated more or less extensively in other tropical countries. Cor- 
responds to the Criollo cultivars. 
Among the collections examined are: 
MEXICO: Veracruz: Colonia San Rafael, flowers and pods, January, 1946; 
forests from Colima to Chiapas, Olivia Converse 74 (UC). Fortufio, Coatzacoalcos 
River, 30-50 m., Ill 1937, "cacao"; tree 15-25 feet tall, crown fairly dense, trunk 
branching from base, inner bark reddish or reddish brown, wood light brown, 
fruit yellow about 8 in. long, 4 in. wide, often cultivated and growing wild in low- 
lands slightly humid or subject to seasonal floods, (LI.) Williams 8457 (F, P). 
Oaxaca: Santa Maria de Chimilapa, I 1927, "wild cacao," MeU 29 (US). 
BRITISH HONDURAS: Stann Creek Valley, Mountain Cow Ridge; tree 5 in. 
diam., 30 III 1940, "wild cacao," Gentle 3292 (MO, NY, US). Middlesex, 200 ft. 
alt.; tall tree of upright habit of growth, generally found growing along river banks, 
fruits dark red, occasional, 19 XI 1929, specimen with fruits, Schipp 178 (UC). 
El Cayo district, Mountain Pine Ridge, 8 V 1931, Bartlett 13108 (F, NY). 
7a(2). Theobroma cacao subsp. cacao forma pentagonum (Bernoulli) Cuatr., 
comb. nov. Figures 5, 6, 20, 23 
Theobroma pentagona Bernoulli, Uebers. Art. Theobroma 6-7, pi. 2, jig. III. 
1869. 
Theobroma cacao subsp. pentagona (Bernoulli) Le6n in Hardy's Cacao Man. 
312. 1960. 
Type.—Bernouilli, op. cit., plate 2} Jig. Ill, i so type, Bernoulli 98 
(GOET). 
Common names,—Cacao lagarto, alligator. 
Uses.—One of the best quality cacaos known. 
Distribution.—Only known in cultivation, being probably a 
mutant cultivar from T. cacao L. originally from Central America; 
mainly cultivated in southern Mexico and Central America, seldom 
in other areas. It is a weaker variety, for which reason it is being 
displaced by more robust and disease resistant varieties, in spite of 
the high quality of its product. 
Collections examined (cultivated): 
514 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
GUATEMALA: Mazatenaiigo, "cult. Marz. 1865 Tkeobroma penlagonum n. 
Bp.?"; specimen with two leaves and one seed, Bernoulli 98 (GOET, isotype) 
Retaluleu, Apr. 1878, Bernoulli & Cario 3151 (GOET). 
COSTA RICA: Turrialba; fruit red, "lagarto" seed white, 7 XI 1961, Cua.tre- 
casas <fc Soria 26540 (US). 
COLOMBIA: El Valle: Finca Esmirna, "lagarto rojo"; seeds white, 16 X 
1961, Cuatrecasas & Barros 26004 (US); ibidem, "lagarto amarillo"; seeds white, 
Cuatrecasas & Barro$ 26005 (US). 
7a(3). Theobroma cacao subsp. cacao forma leiocarpum (Bernoulli) Ducke, 
Rodriguesia 4:274. 1940. Figure 25 
Theobroma leiocarpa Bernoulli, Uebers. Art. Theobroma 6, 7, pi. 2, fig. II. 
1869: (not T. leiocarpum of current authors). 
Theobroma cacao var. leiocarpum Cifcrri, Real Accad. Ital. Mem. Cl. Sci. 
Fis. Mat. e Nat. 4:604. 1933, as to basionym. 
Theobroma cacao subsp. leiocarpum Cuatr. in Macbride, Field Mus. Publ. 
Bot. 13 (3A): 654. 1956, as to basionym. 
Type.—Bernoulli, op. cit., pi. 2, fig. II; isotype, Bernoulli 97 
(GOET), 
Common name.—Cumacaco (Guatemala). 
Uses.—A current cocoa of high quality. 
Distribution.—Atlantic coast of Guatemala and rarely in other 
parts of Central America and southern Mexico. Only known in cul- 
tivation, probably being a mutant cultivar of T. cacao L. The com- 
mercial type known as "Porcelain© Java Criollo" probably represents 
this form. 
Collections examined (cultivated): 
GUATEMALA: Mazatenango, "cult. Marz. 1865, Tkeobroma laeve n. sp.?.," 
Bernoulli 97 (GOET, isotype). Retaluleu, "April 1878, Theobroma laeve Bern.," 
Bernoulli <fc Cario 3152 (GOET). 
Theobroma cacao subsp. cacao forma lacandonense Cuatr., forma no v. 
Figure 25 
Fructus ovoideo-oblongus, acutus, 10-angulatus, 12 cm. longus, 5.3 
cm. diam., pericarpio 5 mm. crasso, epidermide dura, epicarpio molli 
2-3 mm. crasso, mesocarpio cartilagineo, endocarpio molli circa 2-3 
mm. crasso; semina oblonga 20-22 x 14 x 8-10 mm. 
Type in the U.S. National Herbarium, No. 2404633, collected near 
Caribal Lac a nj & ("selva laeandona"), northeast of Chiapas, Mexico, 
in high primary forest (about 50 m. high), 550 m. alt. by F. Miranda 
(No, 9299); half-vine, about 7 m. tall, with very long branches; trunk 
15 cm. diam. The Laeandona Indians call it "cacao." 
Distribution.—Only known from the type locality. It is a very 
interesting variety due to the fact that it is a true wild plant and 
therefore a possible ancestor of the present cultivated cacao. 
CUATRECASAS—CACAO AND ITS ALLIES 515 
MEXICO: Chiapab: Selva Lacandona, Caribal Lacanja, 550 m. alt. in primary 
forest, X 1960, F. Miranda 9299 (US, holotype). 
7b. Theobroma cacao subsp. sphaerocarpum (Chevalier) Cuatr., comb. nov. 
Figures 20, 24, 25 
Theobroma ephaerocarpa Chevalier, Veget. Util. Afr. Trop. Fr. 4:12. 1908. 
Theobroma cacao subsp. leiocarpum sensu Cuatr., excl. basionym T. leiocarpum 
Bernoulli. 
Theobroma leiocarpum sensu Pittier et auctt., excl. basionym T. leiocarpum 
Bernoulli. 
Theobroma cacao var. leiocarpa sensu Ciferri, excl. basionym T. leiocarpum 
Bernoulli. 
Theobroma cacao f. leiocarpum sensu Ducke, excl. basionym T. leiocarpum 
Bernoulli. 
Type.—Sao Tom 6, Chevalier, preserved fruit (P). 
Common names.—Calabacillo, amelonado, Amazonian forastero, 
forastero, etc. (South America). Laranja (Sao Tom6, West Africa). 
Uses.—A currently used cocoa of variable quality; the original type 
gives the lowest quality of the cultivated varieties. 
Distribution.—Native in South America, found spontaneous in 
the Hylaea from the Guianas and middle Amazonia north and west- 
ward to the Andes. Its several cultivars and forms are planted 
throughout the tropics, especially in South America and Africa. Be- 
ing hard and robust trees, and fast-growing plants, they are spreading 
steadily in plantations. 
Some of the collections examined are: 
BRITISH GUIANA: Mataruki River, upper Essequebo; small riparian tree* 
Myers 5829 (K). 
COLOMBIA; Meta: San Jose del Guaviare, forest on left side of the Guaviare 
River, 240 m. elev.; fruits 18 x 7 cm., abundant, apparently spontaneous trees, 
in the lower tree-layer of forest, 14 XI 1939, Oualrecasas 7756 (COL, F, US). 
Sierra de la Macarena, train between Gflejar River and Cafio Guapayita, Cafio 
Yerli, 500-600 m. elev.; slender tree 35 ft. tall, 20-28 XII 1950, Idrobo & Schultes 
784 (COL, US). Ibidem, tree 12 m, tall, 20-28 XII 1950, Idrobo <fc Schultea 940 
(COL, F. US). Ibidem, Cafio Yerli, dense humid forest; shrub 3 m. tall, 25 XI 
1949, Philip son, Idrobo, & Femdndez 1565, 1569 (BM, COL, US). 
AMAZON AS: La Pedrera, Caquet£ River, river level; old trees possibly planted 
long ago and not truly wild on the river banks and islands, pods green, ripening 
yellow, 20 X 1952, Baker & Cope 15 (COL, F, US, TRIN). 
CaqtjbtX: Rio Cagu&n, Cartagena; tree 7-8 m., found distant from river 
bank, white-based pod, smooth and scarcely furrowed, all beans pigmented, 
20IV 1953, Cope & Holliday 105 (COL, TRIN, US). Ibidem; tree 8 m,, numerous 
small fruits smooth, distinctly 10-ridged, somewhat pigmented, very dark purple 
beans, 20 IV 1953, Cope & Holliday 107 (COL, TRIN, US). 
Magdalena: Poponte, in forest of Magdalena valley; tree 30 ft., leaves dark 
green, flowers yellow, fruit red, 16 XII 1924, Cyril Allen 880 (MO). Ibidem; 
fruit green yellow, Allen 881 (MO). 
BRAZIL: Amazonas: Esperan$a, at mouth of Rio Javary, in noninundatable 
rain forest; small tree, flowers whitish, spontaneous "cacao," 25 IX 1942, Ducke 
1095 (IAN, MG, MO, NY, US); ibidem, Ducke 23976 (US). "B." Constant, 
516 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
inundatftble ground; tree; "cacau silvestre," 9 V 1945, Frdes 20882 (IAN, NY); 
ibidem, Frdes 20573 (IAN, NY). Rio Solimoes, on an island below Tabatinga; 
rain forest, slightly inundatable, small tree, flowers white, spontaneous "cacao," 
24 IX 1931, Ducke 23970 (P). 
PERU: Loreto: Rio Ucayali, Laguna de Canchahuaya, 28 X 1898, Huber 
1392 (MG). A specimen collected by Asplund near Tingo Maria (No, 13408) 
may well represent a new species. I did not study this, the material having been 
lent to Dr. Schultes, Botanical Museum, Cambridge. 
The cultivars of Cacao are grouped as follows: 
Fruit elongated and pointed, warty, 10-furrowed, with 5 ridges more 
prominent than the alternate, or only 5, sometimes almost smooth 
and only attenuate not pointed; shell medium thick, easy to cut; 
seeds usually rounded in cross section, white or yellowish white 
inside, and slightly bitter in taste. (Figs. 5, 20, 21, 22, 23.) . . Criollo 
Fruit ellipsoid, rounded at both ends or somewhat narrowed toward 
the apex, rather smooth, with 10 more or less marked furrows; 
shell thick and harder to cut; seeds usually flattened, violet inside, 
bitter in taste. (Some varieties with deep furrows or warty surfaces, 
and pale violet or white beans, being intermediate forms with 
Criollo. (Figs. 5, 6, 20, 22, 24, 25.)   Forastero 
Criollo Morris (1882, pp. 12, 13). This is the type developed 
and propagated since prehistoric times in Central America and 
southern Mexico and which later acquired importance in western 
Venezuela (cacao Caracas). It comprises the best qualities of cocoa, 
but the plants are vulnerable to diseases. Criollo is widely cultivated 
throughout the tropics, the following cultivars being especially 
known by the growers: Venezuela criollo, Nicaragua, Java, Ceylon, 
Samoa, Madagascar, Surinam, and Porcelaine Criollos. See van Hall 
for more information. Among the Criollos has to be included "Alli- 
gator" or "Lagarto" mentioned above as T. cacao f. pentagonum. It 
seems to me that "Porcelaine Criollo" corresponds to T. cacao f. 
leiocarpum. Figs. 5, 20, 21, 22, 23. 
Forastero Morris (1882, pp. 12, 13). This group of cultivars 
was divided by Hart into two: Calabacillo and Forastero, the former 
containing the most typical, shortly ellipsoid, smooth forms. It 
comprises the hardiest, easiest and fastest growing types, but also 
those with lower qualities of cocoa; most Forasteros are an inter- 
breeding product of Calabacillo and Criollo and their segregates. 
Some varieties have developed a good combination of characters. 
They originated and spread in South America and Trinidad, especially 
in the Amazon and Orinoco Valley. The most common known types 
of Forastero are the four listed by van Hall: Angoleta, rather deeply 
ridged and warty, about twice as long as broad, with a wide base; 
Cundeamor, strongly ridged and warty, narrower than half of its 
length, rather acute at the apex, constricted above the base; Amelo- 
nado, oblong, ellipsoid, obtuse, rather smooth with rather shallow 
CUATRECASAS—CACAO AND ITS ALLIES 517 
furrows and not constricted above the base; Calabacillo, shortly 
ellipsoid or almost round, the surface smooth with very shallow 
furrows. Figs. 5, 6, 20, 22, 24, 25. 
The Trinitarios also are Forasteros with features intermediate to 
Criollo, being variable in shape and in seed characters. They prob- 
ably originated by interbreeding in the Venezuelan Orinoco basin, 
from which they were introduced to Trinidad where they acquired 
new genes from old Criollo plantations and developed extensively. 
Later they were brought to Venezuela (receiving the name Trinitario), 
to Ceylon, and more recently to other countries. Trinitarios are 
cocoas with well-balanced conditions of hardness and quality of prod- 
uct. Cacao Nacional of Ecuador is another special Forastero of 
superior quality. (See van Hall, Baker, etc.) 
At the Eighth Inter-American Cacao Conference (1960) an "Inter- 
national Register of Cacao Cultivars" was established, to be organized 
under the chairmanship of Dr. B. G. D. Bartley, of Trinidad. 
Section 4. Telmatocarpus 
Theobroma sect. Telmatocarpus Bernoulli, Uebers. Art. Theobroma 11. 1869. 
Figure 4 
Sect. Bubroma subsect. Telmatocarpus (Bernoulli) Pittier, Rev. Bot. Appl. 
10(110): 779. 1930. 
Petal-lamina lacking. Petal-hood 5-nerved. Staminodes long-cau- 
date, flagelliform, ovate-enlarged at base, flexuose in aestivation. 
Filaments 3-antheriferous. Fruit ovoid, ellipsoid or globose, the 
pericarp thick, costate-nervate-reticulate and lacunose, pilose, or 
tomentulose, the endocarp rigid, thin-woody. Leaves beneath pu- 
berulous or subglabrous. Germination hypogeous. Inflorescences 
axillary on trunk, small, the peduncles solitary or 2 or 3. Main axis 
sympodial with pseudoapical growth; orthotropic shoots from axillary 
buds of the terminal jorquette. Primary branching tern ate. 
Type species: Theobroma microcarpum Mart. 
8. Theobroma gileri Cuatr. Figures 5, 6, 26, 27, 28, 29; Map 7 
Theobroma gileri Cuatr. Rev. Intern. Bot. Appl. 33:562, fig. 1. 1953; Baker, 
Cope & al. (1954) 13, fig. 19; Le6n (1960) 316, 315. fig. 
Type.—Esmeraldas, Ecuador, Manuel Oiler 162 (flowers), 168 
(fruits). 
Small tree up to 14 m. high; growth pseudoapical; trunk 20 cm. in 
diameter, brownish, the bark orange in section, the wood white, 
rather hard; primary branches ternate-verticillate, light brown, warty 
lenticellate, glabrous; terminal branchlets thin, the hornotinous 
stellate-puberulous or pubescent; stipules narrowly subulate, minutely 
stellate-pubescent, 5-8 mm. long, deciduous. 
518 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Map 7.—Geographical distribution of Theobroma microcarpum # and T. gileri Q, the two 
vicariant species (eastern and western of the Andes) of the sect. Telmatocarpus. 
Leaves distichous, thin-coriaceous; petioles 3-5 mm. long, mediocre, 
sub terete, stellate-tomentulose, 3-5 mm. long; blade elliptic-lanceolate, 
narrowed and caudate at apex, asymmetrically rounded at base, 
entire, 6-22.5 cm. long and 1.5-8 cm. broad (usually 7.5-13 x 2-4 cm.) 
including the acumen, this acute, 12-25 mm. long, shining above, 
sparsely stellate-puberulous when young, glabrous or subglabrous 
when adult except for the midrib, sparsely callous-granulate, the 
midrib and secondary nerves conspicuous, with scattered appressed 
stellate hairs beneath, these more copious on the main nerves, but 
the prominent midrib stellate-tomentulose with smaller stellate hairs, 
the 5 or 6 secondary nerves each side filiform, conspicuously prominent, 
arched-ascending, near the margin decurrent and anastomosing, the 
transverse tertiary nerves prominent, the lesser veins prominulous 
loosely reticulate. 
Inflorescences on tubercular growths on trunk and on the branches, 
the cymes few-flowered, often reduced to one flower, the axis very 
short, knotty, bracteate; peduncle erect, filiform, 5-25 mm. long, 
3-bracteolate at apex, the pedicel 7-8 mm. long, somewhat thickened 
toward the apex, scattered pilose; sepals ovate-lanceolate, rather 
thick, reddish or purplish, glabrous inside, greenish or ochraceous 
and stellate-tomentulose outside, minutely tomentulose at margin, 
CUATRECASAS—CACAO AND ITS ALLIES 519 
B 
H 
- 
k 
4 
> 
«a 
* 
I\* 
Figure 27,—a-h, Tkeobroma microcarpum (Ducke 21045): a( b, c, petal from inside) outside, 
and laterally, X 5; d, androecium, X 5; E, stamen, X 10; f, sepal from inside and 
outside, X 2; g, pistil, X 5; h, bud, X 2. i-n, T. gUeri (Giler 162): r, flower on branch; 
j, ovary, X 5; k, androecium, X 5; l, stamen, X 10; m, sepal from inside and outside, 
X 2; n, petal from inside, outside, and laterally, X 5. 
united 1-1.5 mm. at base, about 6 mm. long, 3 mm. broad; petal- 
hoods purplish-red obovate, very narrowed in the lower third, rounded- 
cucullate, muticous or emarginate at apex, with no appendix, sparsely 
covered with slender hairs above, with 5 prominent minutely pilose 
nerves inside, 3-3.2 mm. long, 2.2 mm. broad; androecium purplish 
red, the tube about 1.2 mm. high, minutely puberulous; staminodes 
about 7 mm. long, the base laminar, ovate, 1 mm. long and wide, 
with minute, thick hairs, suddenly narrowed at apex into a subulate, 
flexuous, flagelliform, glabrous appendix about 6 mm. long; filaments 
white, rather thick, glabrous, shortly trifurcate, triantheriferous; 
anthers bilobate, the thecae ellipsoid; ovary ovoid or ellipsoid 5- 
angulate, densely stellate-tomentose; styles glabrous, about 1 mm. 
long, coherent. 
Fruit ovoid or ellipsoid at maturity, 7.5-11 cm. long, 7.5-9 cm. 
broad, brownish green or olivaceous, appressed stellate-tomentose, 
with 5 longitudinal prominulous ribs and irregularly, loosely, reticulate 
prominulous nerves, the intermediate surface shallowly depressed; 
pericarp 1 cm. thick, with a pelliclelike epicarp, a thick carnose, very 
mucilaginous mesocarp, and an inner, 1 mm. thick, coriaceous, 
680-695—64 10 
520 
ligneous, hard endocarp, inside with a sweet, flavorous, ochraceous- 
white pulp surrounding the seeds; seeds ovoid or ovoid-oblong, 2.5-3 
cm. long, 1.8-2 cm. broad, 1.5 cm. thick, usually 20-25 in a pod, the 
Figure 28.—Theobroma gileri (Giler 162, 168): a, leaf, X 1; B, young, ribbed fruit, X 
c, transection of same; d, mature fruit, X E, long, section of same; f, two seeds 
stripped from pulp, X %i o, seed with its pulp, X %. 
CUATRECASAS—CACAO AND ITS ALLIES 521 
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C D 
Figure 29.—A, b, Detail of jndument on the underside of the leaf in: a, Theobroma gileri 
(Giler 162); b, T. microcarpum (Krukoff 1644). c, D, detail of nervation at the under- 
side of the leaf in: c, T. angustifolium (Allen 6259); d, T. subincanum (Cuatr. 7277). 
AX 35, B X 25, C and D X 2. 
522 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
cotyledons white, when fresh condition.1 The juvenile fruits are 
ellipsoid or oblong-ellipsoid, with 5 very strong, protuberant ribs and 
5 less prominent ribs, these sometimes becoming covered by the 
fleshy mesocarp at maturity, but often mature fruits are strongly 
10-ribbed; fruiting peduncle 2-3 cm. long, about 6 mm. in diameter; 
germination hypogeous. 
The fruits of T. gileri are conspicuously larger than those of T. 
microcarpum, its Amazonian vicariant; they are more oblong-ellipsoid 
or more oblong in shape. The young fruits have 10 very prominent 
and thick ribs, 5 of them much stronger than the alternate, and a coarse 
reticulum between the ribs. In a perfect, healthy, mature fruit, the 
surface becomes slightly 5-ribbed and shallowly reticulate-lacunose, 
the fleshy tissue of the mesocarp having filled up the spaces between 
the ribs, as is shown in jig. 28. But often mature fruits keep the 10- 
ribbed, strongly reticulate shape of the young stage, looking very 
similar then to the fruits of T. bicolor. Both kinds and different 
sizes of fruits are usually seen on the same tree. The trees are com- 
monly 8-10 m. high in heavy rain forests. The growth is pseudo- 
terminal, that is to say, by shoots produced above the terminal whorl 
of branches or jorquette. The branching is normally verticillate 
(ternate) but often T. gileri develops adventitious, upright shoots 
which give irregularity to the branching. Fungus diseases may have 
some influence in this. Already in 1953 Patino wrote: "Unfor- 
tunately, in spite of the abundance of trees, this species does not 
seem to offer much commercial prospects, because almost all fruits we 
found were diseased. I sent specimens to Quito to identify the respon- 
sible fungi," and he says further: "Almost all fruits we saw suffer a 
disease which hardens the mucilaginous tissue making it compact and 
hardening the seeds, many are covered by a mashy down, partly or the 
whole." I have seen myself the same, very recently near Villa 
Arteaga. Baker et al. wrote: "Seeds of this species were sent to 
Trinidad but arrived in a decomposed condition. The fungus 
Monilia roreri Ciferri was found infecting fruit of this species." 
Common names.—Chocolate de monte. 
Uses.—Natives used to chew or eat the pulp, which is sweet and 
aromatic. They prepare also a chocolate which is said to have a 
good taste (Patino). 
Distribution.—Restricted to the rain forests of western Colombia 
and northwestern Ecuador. 
COLOMBIA: Antioquia: Villa Arteaga; tree 10-12 m., 15-18 cm. in diameter 
at base, flowers not seen but borne in cushions on trunk and main branches, fruit 
about 9x7 cm., ridged and reticulate, said to be yellow at maturity, jorquette 
1
 Holdrldge reports them purplish when cut, 
CUATRECASAS—CACAO AND ITS ALLIES 523 
symmetrical, 22 VII 1953, Holliday & Bariley T./163 (TRIN, US). Ibidem; tree 
about 10 m., dry fruit 0-9.5 x 7-7.5 cm., no flowers, 24 VII 1923, Holliday & 
Bariley T/167 (TRIN, US). Mutat&, in virgin forest on slope above Villa Arteaga, 
250 m. alt.; small tree, 3 m. tall, 18 cm. DBII, caulifloroua and ramiflorous, sepals 
greenbacked but reddish inside, petals dark blood red, base of central column 
reddish, fruit 10-furrowed, elliptical in long section, about 15 cm. long, seeds 
purplish inside when cut, 23 IX 1959, Holdridge 5133 (US). Ibidem, about 150 
feet alt.; tree 12 m. tall, 6-20 II 1953, SchuUes & Cabrera 18695 (US), Villa 
Arteaga, Las Caucheras, Villa Agraria, rain forest, 200 m. alt.; tree 15 m. tall, 
stem 8 cm. in diam., crown narrow, primary branches ternate, abundant 
inflorescence-cushions (now dry, woody) on the trunk and oldest branches, many 
old fruits on trunk and fallen, and some green ones, hanging, leaves charts ceo us, 
firm, fruits 10 x 7 cm., 10 x 6.5, 11 x 7.5, 11 x 8 cm., 2 X 1961, Cuatrecasas & 
Willard 26167 (COL, US). 
ECUADOR: Esmebaldas: Rio Mira, Guadual, near PiguambI camp, 27 
VII 1953, Giler 162 (holotype, F, 1423965; isotypcs, F, US). Ibidem (fruits), 
Giler 168 (paratype, F). Ibidem, Giler & Patino 164, 165, 166 (H. Cuatr.). 
Probably at Rio Mira; fruit collected by Acosta Soils <6 Giler 12392, 12423 (F). 
Santo Domingo de los Colorados, cultivated from seeds brought from Lita, 
27 IX 1957, Jorge Ledn 4832 (TURRI). 
9. Theobroma microcarpum Mart. Figures 26, 27, 29, 35; Map 7 
Theobroma microcarpum Mart, in Buchn. Repert. 35 : 24. 1830; Linnaea, 
Litt. Bericht. 32. 1831; Bernoulli (1869) 11, pi. 6; Schumann in Mart. 
(1886) 75; Jumelle (1899) 32, fig. 17; De Wildeman (1902) 95; Huber 
(1906a) 273; Ducke (1925) 131; (1940) 271, pi. i, fig. 4; (1953) 14; Chevalier 
(1946) 277; Addison & Tavares (1951) 25, pi. 9, IB, fig. 5; Baker, Cope 
A al. (1954) 12, fig. 11; Le6n (1960) 316, 321, 
Type.—Brazil, Rio Negro, Martins. 
Small tree up to 10-20 m. high; stem 20-30 cm. in diameter; growth 
pseudoapical; crown small; branches much ramose, ternate at least 
when young, brownish rugulose; branchlets thin, the hornotinous 
appressed stellate-tomentose, cinereous or pale ochraceous, becoming 
glabrous in age; stipules narrow, subulate, pubescent, 2.5-4 mm. 
long, soon deciduous. 
Leaves chartaceous, light green; petioles mediocre, sub terete, 
appressed tomentose, when older transversely wrinkled, 4-8 mm. long 
(in very young specimens longer, up to 15 mm.); blades triplinerved, 
elliptic-oblong or obovate-oblong, attenuate near the base, rounded 
or shortly attenuate at apex, abruptly long-acuminate, asymmetri- 
cally rounded at base or sometimes rounded one side, the other 
cuneate, rarely symmetrically cuneate, 6-18.5 cm. long, 2-7 cm. broad, 
including the acumen, this 0.7-2.5 cm. long, the margin entire, with 
scattered minute, simple and stellate hairs above and pubescent costa 
or glabrous throughout, the main nerves filiform, conspicuous, the loose 
reticulum almost obsolete, with sparse, minute, stellate hairs beneath 
or subglabrous, the principal nerves more or less minutely tomentellous 
or glabrate, the costa and the lower pair of lateral ascending nerves 
524 CONTRIBUTIONS PROM THE NATIONAL HERBARIUM 
prominent, the other 2-4 secondary nerves each side thin but 
prominent, ascending, curved near the margin, anastomosing, the 
slender, transverse tertiary nerves prominulous and the minor 
reticulate veins less prominulous, but conspicuous. 
Inflorescences axillary or extra-axillary on young leafy branchlets, 
the cymose clusters extremely small, bearing 1-3 flowers, the woody 
primary branches short, knotty, bearing ovate, amplectant bracts, 
0.6 mm. long and wide; peduncles rather thick, 0.5-1 mm. long, 
tomentulose, with 3 subulate, deciduous bracteoles at apex, these 
0.6-1 mm. long; pedicels 0.5-1 mm. long, moderately thick, tomentu- 
lose; buds ovoid, acute, 4.5 mm. long, 3 mm. broad, with 5 whitish, 
minutely tomentulose lines and scattered, minute stellate hairs. 
Sepals thick-membranaceous, lanceolate, acute, united about 1 mm. 
at base, 5-6 mm. long, 2 mm. broad, with scattered, minute, stellate 
hairs outside, minutely, whitish tomentulose at margin, with con- 
spicuous midrib and sparse, mediocre, flexuous, glandular hairs 
inside; petal-hoods pale brown, thick-membranaceous, glabrous, 
oblong-obovate, rounded-cucullate and acuminate at apex, the acumen 
triangular, shortly bidentate, exappendiculate, the 5 nerves rather 
thick, prominent and minutely papillose inside, often confluent into 
pairs near the base; androecium red or reddish, the tube about 1.5 
mm. high, 10-furrowed, minutely pilose; staminodes minutely pilose 
with a rather thick, oblong-ovate concave base, this 2.5 mm. long and 
1 mm. broad with fine flexuose hairs inside, topped by a subulate, 
flexuose tail 4-5 mm. long; filaments rather thick, 1 mm. long, shortly 
3-furcate, triantheriferous; cells of anthers ellipsoid, about 0.4 mm. 
long; ovary 1,5-1.8 mm. long, pyriform, glabrous or sparsely, minutely 
stellate-pilose, sharply 10-furrowed-costate; styles 5, united in the lower 
fourth, rather thick, acute, glabrous. 
Young fruit ellipsoid (±3 cm, long), strongly 10-ribbed, with 5 very 
thick and prominent, dorsal, shortly pinnate costae, and 5 smaller, 
commissural ones; mature fruit 6.5-7 (-9) cm. long, 5.5-6.5 cm. 
broad, green yellowish, puberulous, ellipsoid-globose, conspicuously 
10-ribbed, the surface between the ribs shallowly alveolate, the 
exocarp thick, carnose, padding the hollows between the lignose 
underlayer, when dry the ribs and the lignose reticulum become ex- 
tremely marked and prominent; seeds more or less compressed, ovoid, 
12-14 mm. long, 18-20 mm. broad, and 11-12 mm. thick; fruiting pe- 
duncle thick, robust, 4-8 mm. long and broad; cotyledons hypogeous 
at germination. 
Common names.—Cacauf, cacaurana, cacao rana, cacau bravo, 
cabega de urubu (Brazil). Cacao de monte (Colombia). Me-tr6- 
ree-moo-ee (Karihona, Upper Apaporis); b<5o-e (Mirana, CaquetA 
River) (Angl-Col. Cocoa Exp., Baker, 1952). 
CUATRECASAS—CACAO AND ITS ALLIES 525 
Uses.—None recorded on the use of its sweet scentless pulp or the 
seeds. 
Distribution.—In the southern and western upper Amazon 
Brasilian region Rios Solimoes, Yapurd, Pur (is, Madeira, Tap aj 6s, 
and western Colombia on Caquetd River. (Baker & Cope.) The 
eastern known limit according to Ducke is Rio Tapaj 6s. The specimens 
around Bel6m are cultivated. It is frequent in its area and may 
become abundant in some places as a significant element of the shad- 
owy under layer of rain forests on elevated ground and in moder- 
ately inundatable alluvial lands. 
COLOMBIA: Amazon as: Rfo Caquetd, La Pedrera, river level; tall tree 15- 
20 m., 30 cm. in diam. at base, extensive branch system, jorquettes of seedlings 3- 
branched, growth continuing from above, flowers small, petals without ligules, 
fruit abnormal due to attack by Marasmius perniciosus, 5 X 1952, Baker & Cope 
28 (COL, K, TRIN, US). Ibidem; tall tree 15-20 m., native in forest on the river- 
bank, 7 X 1952, Baker & Cope 29 (COL, F, K, TRIN, US). Ibidem; tree 30 
feet, 9 inches in diameter, on floodbank, 5 X 1952, Schultes & I. Cabrera 17780 
(AMES, US). Rfo CaquetA, Remolino; leaves only from small seedling tree 
2.5 m., typical Theobroma habit, jorquettes arising symmetrically, 2 V 1953, 
Cope & Holliday T/125 (COL, TRIN, US). 
BRAZIL: Amazonas: In sylvis ad Costa de Ubicuna et de Camaroeoari, fluv. 
Solimoes, prov. Rio Negro ("Dr. Martius Iter Brasiliensis, 321"), Martina Observ. 
2890, [884] (M, lectotype, photo F, M. 19643). Ibidem, Martius Observ. 2890, 
[885, 886] (M, isotypes). Lower Rio Yapurtf,, Jubar& matta, 15 IX 1904, Ducke 
6773 (BM, MG). Basin of Rio Solimoes, Municipality S&o Paulo de Olivenga, 
near Falmares; tree 60 ft. high, trunk 7 inches in diam., terra firma, high land, 
IIIX-26 X 1936, Krukoff 8280 (A, BM, F, G, GB, K, LE, MICH, MO, P, S, U, US, 
USD A). Ibidem; mata, caatinga, "cacau bravo," arvore pequefia, 19 IV 1945, 
Frdes 20750 (IAN, USDA), 34814 (IAN). Basin Rio Madeira, Municipality Hu- 
mayta, near Livramento, on Rio Livramento on varzea land; tree 50 feet high, 
"cabesa de Urubd," 12 X-6 XI 1934, Krukoff 6592 (A, BM, F, G, K, LE, MICH, 
MO, S, U, US, USDA, WU). Ibidem, Municipality Humayta, near Tres Casas, 
on restinga alta; tree 60 ft. high, 14IX-11 X 1934, Krukoff 6203 (A, F, G, K, LE, 
MO, S, U, US, USDA, Y). Rio Purus, Bom Lugar; "cacao rana," II 1904, Goeldi 
4228 (BM, G, MG), Camatian; high forest lowland, border of creek; tree 7 m. 
high, 25 cm diam., 24 I 1949, Frdes 23963 (IAN, US). 
Guapoke: Porto velho, Entrada de Redagan, Km. 8, Viana, mata derrum- 
bada, terra firme; arvore pequena, 31 V 1952, Black, Cordeiro, & Francisco 52- 
14649 (IAN, UC). 
Mato Grosso: Machado River region, source of the Jatuarana River; tree 
3 feet high in terra firma, "cabeca de urubu," XII1931, Krukoff 1644 (A, BM, F, G, 
K, MICH, MO, P, S, U, UC). 
PahX: BeI6m, Jardim Botanico do Museu Goeldi; medium tree, cultivated, 
11 VIII 1942, Archer 7551 (F, IAN, K, USDA). Beldm, Horto Botanico Par6 
(cultum proven, Rio Purus, Bom Lugar anno 1904), 25 V1906, Hither 7081 (G, MG). 
Ibidem; arbor parva floribus rubcscentibus, 4 II 1926, Ducke 21045 (G, GH, K, S, 
U, US). Ibidem; arbor parva, floribus pallide brunnescentibus, "cacaohy," 
IX1936, Ducke 283 (A, F, K, MO, S, US). Ibidem, 21 VII1944, F. C. Camargo 8 
(IAN). Ibidem, 23 XI 1945, Fires & Black 742 (IAN). Rio Guama, near Bel£m, 
"cacao bravo," IV 1929, Dahlgren & Sella 10 (F, GH). Rio Tapaj os, Cachoeira 
do Mangabal, beira de assahyzal, 7 IX 1916, Ducke 16466 (BM, G, MG, P, US). 
526 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
COSTA RICA: (cult.) Limon, La Lola, experimental station IICA; tree 4 m. 
tall, narrow crown, eight years old, first flowers this year, 7 XI 1961, Cuatreeasas 
tfe Paredes 26538 (US). 
Section 5. Glossopetalum 
Theobroma scctio Glossopetalum Bernoulli, Uebcrs. Art. Theobroma 11. 1869 
Figures 1, 3, 4; Map 2 
Sect. Bubroma Schum. in Engl. & Prantl. Nat. Pflanzenfam. 3(6) :89. 1890. 
Sect. Bubroma subsect. Glossopetalum (Bernoulli) Pittier, Rev. Bot. Appl. 10 
(110): 779. 1930. 
Petal-laminas obovate, spatulate or trapezoid, stipitate. Petal- 
hoods 7-nerved. Staminodes laminar, petaloid, obovate or broadly 
lanceolate, curved-reflexed covering the hoods in aestivation, erect or 
reflexed in anthesis. Filaments 3-antheriferous. Fruit ellipsoid or 
oblong, smooth or more or less angulate or tuberculate, the epicarp 
hard, woody, with a tomentose epidermis. Cotyledons hypogeous 
at germination. Leaves beneath reticulate-nerved, stellate-tomen- 
tose. Inflorescences on the main trunk or on the branches. Main 
axis sympodial with pseudoapical growth; orthotropic shoots from 
axillary buds of the terminal jorquette. Primary branching ternate. 
Type species.—Theobroma grandiflorum Schumann. 
10. Theobroma angustifolium Mogi no & Sess6 
Figures 6, 25, 29, 30, 31, 37; Map 8 
Theobroma angustifolium Mogifio et Sess6 ex DC. Prodr. 1: 484. 1824; Icon. 
Fl. Mexicans ex DC. pi. 118; Bernoulli (1869) 12, pi. 6; Schumann in Mart. 
(1886) 77; Donn. Smith in Pittier, Prim. Fl. Costar. 96. 1898; Preuss 
(1901) 255, pi £,0;De Wilde man (1902) 96, figs. IB,IS; Standley (1923) 
808; Standley (1937) 687; Chevalier (1946) 282; Standley & Steyermark 
(1949) 421; Holdridge (1950a) 3; Allen (1956) 342, pi. 26; Le6n (1960) 
318, 315, fig. 
Type.—Ses$6 et Mogino, Plantae Novae Hispaniae, Herbarium 
Florae Mexicanae. 
Tree 8-26 m. high; trunk up to 30 cm. in diameter, with smooth 
bark and whitish wood; growth pseudoapical; primary branches ternate; 
lower branches horizontal, the higher ascending; branchlets when 
young green ochraceous, densely and moderately appressed tomentose, 
with very minute, fine, translucid-white stellate hairs intermixed with 
other mediocre, fulvous or ochraceous, somewhat thicker, stellate 
hairs, when older more or less glabrate, grayish, rugulose; stipules 
lanceolate-subulate, acute, broadened at base, above sparsely, below 
densely stellate-tomentose, 5-7 (-15) mm. long, about 1 (-2) mm. 
broad, deciduous. 
Leaves distichous, thin-coriaceous, rather flexible; petioles moder- 
ately thick, densely subappressed tomentose, 6-10 mm. long; blades 
subobovate-oblong, elliptic-oblong, or oblanceolate, slightly narrowed 
CUATRECASAS—CACAO AND ITS ALLIES 527 
Map 8.—Geographical distribution of Thtobroma angustifolium # and 7". Jtmiarum Q. 
to the obtuse and slightly asymmetrical base, attenuate and acuminate 
at apex, entire or at the upper part slightly sinuate-dentate, 9-25 cm. 
long and 3-9 cm, broad, including the 1-2 cm. long and 3-5 mm. wide 
acumen, green above, when adult smooth, glabrous or with few hairs 
scattered on the costa, this depressed, filiform, the secondary and 
tertiary nerves little conspicuous, light greenish or cinereous beneath, 
appressed tomentose, heteortrichous, the surface covered with a dense 
layer of white, minute, stellate hairs, and additional, more or less 
copious, larger, ochraceous, stellate hairs with longer rays on the main 
nerves, the costa very prominent, the 6-8 secondary nerves on each 
side thinner and prominent, ascending, near the margin decurrent, 
becoming slender, vanishing, the transverse tertiary nerves, thin, 
prominulous, 2-5 mm. from each other, the minute reticulum conspic- 
uous. 
Inflorescences usually abundant on the branchlets, axillary or 
extra-axillary, the cymes strongly reduced to a few extremely short 
1-3-flowered branchlets; peduncles 0.4-1 mm. long, 3-bracteolate; 
pedicels erect, rigid, mediocre, densely ochraceous or ferruginous, 
ebracteate, 5-10 mm. long; bracteoles very minute (1-0.5 mm. long), 
linear, deciduous; buds globose, 7-8 mm. broad, densely ochraceous 
tomentose; calyx 8-9 mm. long, reflexed in an thesis, all the sepals 
united to 3-4 mm. into a cupular base, in the upper part two united 
528 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
■<■1 
i * ■ 
B 
r "* I / .'v i-'V'jltf'V ■ f - *«?" 
;n-\ \ f viX-'X- - > -% 
c D 
Figure 30.—Detail of indument on the underside of leaf in: a, T. angustifoiium (Allen 
6259); b, T. cinnolinae (Cuatr. 14897); c, detail of venation in T. stipu'alum (Cuatr. 
21339); d, indument in T. stipulatum (Cuatr. 21339), A, B, and D X 30, C X 2. 
Figure 31.—a-f, Theobroma nemorale (Patiiio 116): a, d, petal from inside and laterally, 
X 5; c, androecitim, X 5; d, stamen, X 10; e, gynoecium, X 5; r, sepal from inside 
and outside, X 2. g-m, T. angustifoiium (Allen 6341): G, h, petal from inside and 
laterally, X 5; I, androecium, X 5; j, stamen, X 10; k, gynoecium, X 5; L, bud, X 2j 
CUATRECASAS—CACAO AND ITS ALLIES 529 
[Figure 31] 
m, sepal from inside and outside, X 2. n, o, T. nemorale (Cuatr. 26007): n, the three 
bracteoles covering the opening flower, X 2; o, pe dice lied bud surrounded by a 
bracteole, the other two removed, p, T. ckocoense, st ami node, X 5 (Cuatr. 26074), 
CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
[Figure 32] 
CUATRECASAS—CACAO AND ITS ALLIES 531 
by pairs forming 3 unequal lobes, two of the lobes twice as broad as 
the third, ovate, rather obtuse, scarcely puberulous and with minute, 
thick, oblong glandular hairs at the base inside, tomentose, greenish 
ochraceous or ferruginous outside, each single sepal 4-5 mm. wide. 
Petal-hoods thick-membranaceous, yellowish, broadly obovate, 
rounded-cucullate at apex, 7-nerved with very sparse, thin hairs 
outside, about 5 mm. long, 4 mm. broad; petal-lamina pedicellate, 
yellow, thick-membranaceous, 5-nerved and finely veined, subobovate- 
spatulate, emarginate at apex with 2 ovate or rounded lobes, atten- 
uate towards the base, glabrous, about 5.5 mm. long, 3 mm. broad, 
the pedicel linear, 3-nerved, attenuate downward, about 4-5 mm. 
long, 1 mm. wide; androecium tube 2.5-3 mm. long, thick; staminodes 
laminar, thick-membranaceous, sulphur yellow, glabrous, oblong- 
obovate, rounded or subspatulate at apex, with marked venation, 
8-11 mm. long, 4.2-5 mm. broad, at base 1.2 mm. broad, when in 
bud curved-reflexed, in an thesis erect; filaments glabrous, thick, 
curved, 2 mm. long, enlarged and shortly 3-furcate, 3-antheriferous; 
anther cells globose; ovary obovoid-oblong, about 1.5 mm. long, 
5-sulcate, densely stellate-tomentose; styles 2.5 mm. long, united in a 
rigid erect column 1.5 mm. long, ending in 5 slender branches 1 mm. 
long. 
Fruit unequally oblong-ellipsoid or ovoid-ellipsoid, more or less 
pentagonal, slightly attenuate at apex, umbilicate and 5-costate at 
base, very irregularly tuberculate-rugose, densely brown tomentose, 
the epicarp hard, ligneous, about 1.5 mm. thick, the mesocarp plus 
endocarp carnose 5-6 mm. thick, the pulp enveloping the seeds thick, 
juicy, aromatic, edible, 10-18 cm. long, 6-9 cm. broad; seeds 5-7 in 
each fruit compartment, compressed oblong-ovoid, 26-32 mm. long, 
16-19 mm. broad, and 14-16 mm. thick, the cotyledons white; ger- 
mination hypogeous. 
The leaves of T. angustifolium are similar to those of T. nemorale, 
but they are narrower, rather lanceolate, and the larger hairs of the 
double indument beneath are longer with much finer, longer rays 
than those of T. nemorale. Paul Allen (1956) writes about this tree: 
"The young branches, petioles, and veins of the lower leaf surface are 
covered with a rather scurfy pale-tan torncnturn. The relatively 
Figure 32.—a-g, Thtobroma Jlipulatum (Cuatr. 21339): a, b, petal from inside and later- 
ally, X 5; c, androecium, X S;d, stamen, X 10; E, gynoecium, X 5; r, sepal from inside 
and outside, X 2; o, bud, X 2. h-n, 7*. simiarum (Tonduz 7313): h, i, petal from 
inside and laterally, X 5; j, androecium, X 5; k, stamen, X 10; l, gynoecium, X 5; 
m, sepal from inside and outside, X 2; N, bud supported by pedicel and 3-bracteoIate 
peduncle, X 2. o-u, T. cirmolincu (Cuatr. 14897): o, p, petal from inside and lat- 
erally, X 5; Q, androecium, X 5; r, stamen, X 10; s, gynoecium, X 5;t, sepal from 
inside and outside, X 2; u, pedunculate bud, X 2. 
532 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
■■ ' 
fif i , / . ..: y \sr* 
?, '.VPr/% 
1 
B 
&* 
t 
<Tv. 
"A ■■■■ h "> .. */-.H' 
<y> 
u 
■■'■?■:» 
: ■■* \ -:s* 
jiv? 
: >■ ::-/,■ \U,- ■ ; jC 
;:: y ■/. -,n ..,, V.; r,_^>:'. 
i." j : ■ u,.. v ' .■' i 
V/.■■■■■■ --■ . ■ fr"i- 
v ■ ■=- f ■ * . ■ « j 
, K ->: * ■ t 
'. : , \ . .; 
■ K.J-* 
sr 
w 
[Figure 33J 
CUATRECASAS—CACAO AND ITS ALLIES 533 
large, bright-orange flowers are produced in great profusion in several 
successive flowerings from November until February from the axils 
of the slender, younger branches, and are followed in August and 
September by the large, woody, brown-tomentose, cacaolike pen- 
dulous pods which are from 4" to about 7" in length." 
Common names.—Cacao de mico, cacao silvestre (Costa Rica), 
cushta, cacao de la India (Salvador), cacao silvestre (Mexico). 
Cacao de mico, cacao meco, coca mono (Nicaragua), sor<5 (Bribi 
Indians). 
Uses,'—It was stated by Standley (1923) that T. angustijolium was 
much planted in southern Mexico, especially in Chiapas as a source 
of commercial cacao and that the famous Soconusco cacao was from 
this species- This statement is very doubtful, for the seeds of T. 
angustijolium are considered at present in that area as of inferior 
quality without commercial value. 
Distribution.—This species is often planted in Central America 
and southern Mexico. It is certainly native in the lowland forests of 
the Pacific range of Costa Rica (Allen) and nearby Central American 
countries (Holdridge). Standley and Steyermark (1949, 422) say 
that the native region of this cacao is unknown, but Tonduz already 
in 1891 cited it from the forests of Terraba; Le<5n (No. 937) writes: 
"Important tree in the regional forests," and Allen says that "it is 
locally frequent in lowland forests throughout the area [Golfo Dulce]." 
MEXICO: Direcidn de Estudios Biol6gicos, Ord. 34-G823 (MEXU). Her- 
barium Sess6 & Mo$ifio in M, "18-1, Theobroma Simiarum N. Ic," SessS, 
Castillo, dt Maldonado 3618 (hoiotype, MA; isotype, F, Photo F. M\ 48411), 
Swat & Mosifio s.n. (BM, probable isotype). Copy of the Sess6 & Mogifio drawing 
at G (Negative F. M. 30527), plate 112 of the DC. published series. 
GUATEMALA (cult.): Retaluleu, April, 1877, Bernoulli & Cario 3188 (GOET, 
K, S). Mazatenango, III 1865, cult., Bernoulli 95 (NY, BR). Region of Plata- 
nares, between Taxi so o and Guazacap&n (Dept. Santa Rosa), 220 m. alt., wet 
forested quebrada; small tree escaped here, 3 XII 1940, Standley 79068 (F, US). 
SALVADOR (cult.): Sonsonate, "cushta," 1922, Calderdn 630 (GH, NY, US). 
Vicinity of Sonsonate, 220-300 m. alt.; tree 20-30 feet, very dense and narrow 
crown, flowers on branches, fruit brown, the pulp edible with very aromatic odor, 
seeds give chocolate, grown here only in finca, "cushta," "cacao de la India," 
Standley 22317 (GH, MO, NY, US). 
Figure 33.—a-h, Theobroma grandtflorum (Ducke 598): A, b, petal from inside and 
laterally, X 5; c, androecium, X 5; d, stamen, X 10; E, gynoecium, X 5; r, styles, 
X S; G, sepals from inside, X 2; h, sepals from outside, X 2. i-o, 7". obovatum (Ducke 
265): i, j, petal from inside and laterally, X 5; k, androecium, X 5; l, stamen, X 10; 
m, gynoecium, X 5;n, sepal from inside and outside, X 2; o, bud supported by bracte- 
olate peduncle and pedicel, X 2. p-w, T. subincanum (Baker & Cope 32 and Holliday 
43): p, q, petal, from inside and laterally, X 5; x, androecium, X 5; a, stamen, X 10; 
T, bud, X 2; u, sepal from inside and outside, X 2; v, initiation of fruit, X 5; w, ovary, 
X S. 
534 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
NICARAGUA: Quezalguaque, Dept. of Le6n, Baker 2102 (A, AMES, C, GH, 
G, K, L, MICH, MO, NY, XT, UC, US, WU). Chichigalpa, "cacao de mico," 
II 1900, Preuss 1381 (UC). Beldo, "cacao meco," "coca mono," or "monkey 
cocoa," 28 VI 1893, Hart 5381 (K, NY, U). 
COSTA RICA: Guanacaste: Upper portion of cafi6n of Rfo San Jos6, 460-480 
m., 12,13 II1930, Dodge & Thomas 6399 (F, GH, MICH, MO, UC, US). Nicoya, 
300 m. alt., VI 1949, Lopez B.n. (F, TURRI). Ibidem, Pittier s.n, (US). Ho- 
jancha de Nicoya, 20 ra. drbol importante en la selva de la region, "cacao de mico," 
29 I 1942, Le6n 937 (F). Perico, Nicoya, 100 m., I 1954, Le6n 4267 (TURRI). 
Borde du Rto Zurqufn, 3 1894, Pitiier & Durand 8536 (P). 
Limon: Cienaguita, near Puerto Lim6n, 10 m. alt.; small tree with ap- 
pressed crown, flowers sulphur yellow, VII 1901, Pittier 16142 (G, US, WU). 
La Lola, I.I.A.C.A. experimental station, about 20 m. alt.; tree erect, trunk about 
25 cm. diam. at base; pseudoapical growth; primary branches ternate, spreading, 
persistent; leaves thin-coriaceous but firm, light green and somewhat glaucous and 
cinereous beneath; young fruits thickly tomentose, axillary flowers abundant, now 
dry, 6 XI 1961, J. Cuatrecasas & Paredes 26537 (US). 
PuNTARENAS: Dans le for£t k Terraba, 260 m. alt., II 1891, Tonduz 4074 
(US). Ibidem; "cacao de mico," II 1891, Pittier & Tonduz 4074 (BR, G). 
Boruca, Diquis Valley, 1891, Piltier s.n. (US). Tinoco Station, fairly frequent in 
swampy forest; tree 80 ft., fruits pendulous, produced from the ends of branches, 
"cacao de mico," 13 VII 1951, Allen 6259 (BH, MO, US). Lowland forest near 
Palmar Norte; tree 45 feet, flowers bright orange, Allen 6341 (BH, F, MO, US). 
Llanuras de Corredor (Golfo Dulce), III 1897, Pittier 11112 (G). 
PANAMA: Progreso (Chiriqui); small tree 30 feet, 6 inches in diam,, with a 
fruit tike wild cacao except that the husk is smooth like a potato skin, big seeds 
with white meat inside, 1927, Cooper & Slater 242 (F, GH, NY, US, Y). Comarca 
del Bard, Puerto Armuelles, United Fruit Company farms between Canasco and 
Cocos, mostly cutover land with some of the original trees still standing, about 
100 feet alt.; tree, fruit resembling a cacao pod; leaves pale bluish, green, beneath, 
17 VI 1957, Stern & Chambers 140 (MO, US, Y). 
TRINIDAD: Royal Botanic Gardens, Port-of-Spain, L. H. Bailey, s.n. (BH), 
Ibidem; small tree, flowers orange colour, 8 IX 1918 (originally from Guatemala), 
Broadway 8935 (BM, BR, MO, S). Ibidem; tree 8 m., flowers orange yellow, 
10 VII 1953, R. E. D. Baker s.n. (TRIN). Government House Gardens, 12 III 
1929, Williams 12121 (TRIN). St. Augustine, Imperial College of Tropical Agri- 
culture; tree 3-4 m. alt., lower branches horizontal, upper ones ascending, bark 
granulose, lenticellate, more or less cleft, yellowish brown, abundant, hanging, 
very rugose dried fruits, 1 IX 1961, Cuatrecasas & Cope 25789 (US). Ibidem; 
tree about 8 m, (9 years old), trunk 20 cm. in diam. at base, primary branches 
ternate from near the base, leaves thin-coriaceous, yellowish green, shining above, 
greenish cinereous beneath, 1 IX 1961, Cuatrecasas & Cope 27591 (US). 
11, Theobroma cirmolioac Cuatr. 
Figures 3, 4, 30, 32, 34, 36, 37; Map 9; Plate 7 
Theobroma cirmolinae Cuatr. Notas Fl. Colomb. VI: 5, fig. 1-5. 1944; 
Rev. Acad. Colomb. Cienc. 6:32, fig. 1-5. 1944; Llano (1947) 34, 
pi. 14; Baker, Cope & al. (1954) 13, fig. 22; Le6n (1960) 320, 317, fig. 
Type.—El Valle, Colombia, Cuatrecasas 14897 
Medium-sized or large tree up to 20 m. high; growth pseudoapical; 
trunk up to 40 cm. in diameter, branched in the upper third, the bark 
dark grayish, somewhat rimose-scaly, under the periderm brown or 
CUATRECASAS—CACAO AND ITS ALLIES 535 
Map 9.—Geographical distribution of Theobroma Jtipulatum #, T. cirmolinae 0» T. 
ckocoeme 0, T. mammosum 0, T, sinuosum A, and 7*. canumanense fl. 
rufous, the wood ochraceous, the hardwood ochraceous reddish, very 
hard; branches gray or brownish gray, the primary ternate, the 
terminal leafy branchlets tawny or ferruginous, appressed stellate- 
tomentose; gum resin flowing easily from bark and wood; stipules 
large, persistent, subcoriaceous, oblong-lanceolate, subacute, ochra- 
ceous-tomentose, 12-22 mm. long, 3-5 mm. broad at base. 
Leaves of the young branchlets (smaller than in adult) thin- 
chartaceouSj green-ochraceous with scattered stellate hairs above, 
green ochraceous or grayish green, appressed stellate-tomentose 
beneath; adult leaves large, thick-coriaceous, rigid, shortly petiolate; 
petioles very strong, thick, sub terete, appressed ochraceous-tomentose, 
1-2 cm. long, 6-8 mm. broad; blades oblong-elliptic or ovate-elliptic, 
slightly asymmetrical, rounded, cordate, or sharply emarginate at 
base, little attenuate, rounded or very obtuse and shortly acuminate 
at apex, entire or very slightly sinuate and flat at margin, 26-54 cm. 
long, 14-30 cm. broad, ochraceous green above, pale brown when 
dry, apparently glabrous but with scattered, appressed stellate 
hairs, these more copious on the main nerves, the costa and the 
secondary nerves filiform and depressed, the minor venation less 
noticeable, velvety-tomen tose beneath, the surface rosy-glaucous, 
the veins somewhat more ferruginous or rufescent, the costa very 
thick and prominent, the 12-14 secondary nerves on each side very 
080-695—64 11 
536 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
••••><* 
Figure 34.—Thtobroma cirmolinae, flowering and fruiting trunk, X V%. 
CUATRECASAS—CACAO AND ITS ALLIES 537 
Figure 35.—Fruits of Theobroma, X a, bicolor (Llano s.n.); b, stipulation (Cuatr. 
21339); c, subincanum (Little 9544); d, microcarpum (Archer 1551); e, obovatum (KJug 
2983). 
prominent, subspreading, near the margin arched, decurrent, anasto- 
mosing, the transverse tertiary nerves prominent, those of the fourth 
rank also prominent, reticulate-anastomosing, the lesser veins forming 
a minute, prominulous reticulum, the minor reticulum and areoles 
covered by a dense tomentum of intricate, white, sericeous, minute 
538 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
stellate hairs, the major nerves with only scattered minute, slightly 
larger hairs and abundant, minute reddish, callose warts; base of lamina 
7-nerved, the 4 lower nerves small, of lower degree. 
Fertile branches perennial on main trunk and big branches, short, 
ligneous, tuberculate, prolific in flowering, the short, intricate, cin- 
Figure 36.—Fruits of Thtobroma, X 'a :^A, simiarum (Cuatr. 26515A); b, simiarum (Cuatr. 
26536); c, ckocoense (Patifio 115); d, cirmolinae (Cuatr. 15336); E, grandiflorum (Cuatr. 
25780T); f, mammosum (Cuatr. 26535). 
CUATRECASAS—CACAO AND ITS ALLIES 539 
Figure 37*—Leaves of Theobroma: stipulatum (Cuatr. 21339); s, stipulatum, stipules and 
base of leaf beneath (21339); c, citmolinae (Cuatr. 15336); D, angustifolium (Allen 
6259); E, angustifolium (Stand, 22317)j showing the stipules* All leaves X Y%X 54- 
540 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
cinnate branches forming cushions up to 12 cm. broad on the trunk; 
sympodial branchlets angulate, ramose, bracteolate, forming crowded 
panicles up to 8 cm. long, appressed stellate-tomentose or glabrate 
when old; bracts 1,5-3 mm. long, 1,5-2.5 mm. broad, subcoriaceous, 
persistent, subamplectant, triangular-ovate, tomentose; peduncles 
elongate in anthesis, erect, rather thin, ferruginous tomentose, 5-8 
mm. long, 3-bracteolate at apex; bracteoles linear, subacute, 3-6 mm. 
long, 1-1.5 mm. wide; pedicels in an thesis 15-20 mm. long, thin, 
erect, tomentose, ebracteate; buds globose, with 5 commissural ribs, 
densely ferruginous tomentose. 
Sepals thick, carnose, ovate-triangular, acute, united in the lower 
third or fourth, soon umbilicate-reflexed, with the five free lobes 
spreading, 10-15 mm. long, 6-7 mm. broad, pale yellow inside, 
glabrous, except for the base, with minute, oblong, glandular hairs 
at their insertion, yellowish, thick-tomentose outside with ochraceous 
or tawny stellate hairs, the margin minutely cinereous-tomentose. 
Petal-hoods yellow, thick-membranaceous, elliptic-obovate, rounded- 
cucullate at apex, glabrous, with 7 prominent nerves inside, 6-7 mm. 
long, 4-5 mm. broad; petal-laminae thick, carnose, glabrous, sub- 
triangular-spatula te, slightly 3-undulate or emarginate at apex, about 
3.5 mm. long, 2.2 mm. broad, tapering to a pedicel at base; pedicel 
about 4 mm. long, 0.6 mm. wide, bidentate at the joint. 
Androecium tube thick, 2-2.2 mm. high; staminodes laminar, 
rather thick, oblong-spatulate, slightly broadened towards the end, 
retuse at apex, erect, glabrous, sulphur yellow, often somewhat 
reddish near the base, 10-11 mm. long, 4-5 mm. broad, at base 1.5 mm. 
wide; filaments robust, glabrous, 2.2-3 mm. long, shortly 3-furcate, 
3-antheriferous, rarely 4-furcate, 4-antheriferous; ovary ovoid, 1.6-2 
mm. long, 1.8 mm. broad, 5-costate, tomentose; styles 2 mm. long, 
connivent, free up to near the base. 
Fruiting peduncle robust, 3-A cm. long, 8-10 mm. thick, articulate; 
young fruits large, fusiform, prismatic with 5 obtuse, prominent ribs 
corresponding to the loculi and 5 others, more or less marked, alternate 
commissural ones, the base umbilicate, the apex attenuate and obtuse; 
ripe fruit 25-35 cm. long, 10-12 cm. wide, ellipsoid-oblong or obovoid- 
oblong, very little narrowed to the umbilicate base, attenuate to the 
obtuse apex, the surface obtusely pentagonal, with rounded ridges, 
brown or ferruginous, stellate-tomentose; pericarp about 1-1.5 cm. 
thick; epicarp 2 mm. thick, very hard, ligneous, the mesocarp and 
endocarp carnose, becoming hard and coriaceous when dry; covering 
of the seeds fibrous, pulpy, yellowish white and flavorous; seeds striped 
from the more or less compressed pulp, ovoid, the testa brown, the 
outer tegument light brown, the inner one dark brown, the cotyle- 
CUATEECASAS—CACAO AND ITS ALLIES 541 
dons reddish, 20-24 mm. long, 15-19 mm. broad, and 9-16 mm. thick; 
germination hypogeous. 
Common names,—Bacao, cacao de monte, cacao indio. 
Uses.—No special uses known besides occasional preparation of 
chocolate by the natives. This species is the one which grows at the 
highest altitudes; it should be tried as a grafting base, especially in the 
coldest zone of cacao production. 
Distribution.—Only known from the Pacific slopes of the Andes 
in Colombia in the Department of El Valle, between 800 and 1300 
meters altitude. 
COLOMBIA: El Valle: Western slope of western Cordillera, valley of 
Rio Digua, Piedra de Moler, rain forests, 900-1180 m. alt-; tree 20 m., trunk 40 em. 
in diam., adult leaves thick-coriaceous, green above, yellowish green beneath, 
sepals ferruginous-green, petals yellow, staminodcs bright yellow, their bases 
reddish, bark and central wood with resine, "bacao," 19 VIII 1943, OuatrecaM8 
14897 (holotype VALLE; isotypes, F, Y). Valley of Rio Dfgua, La Elsa, forests 
1000-1200 m. alt.; tree 12 m., branched in the upper part, bark dark gray, almost 
smooth, flowers yellow, 9 XI 1943, Cuairecasas 15336 (F, VALLE, Y, paratypes). 
La Elsa, about 800 m. alt.; tree 12-15 m., 30-35 cm. in diam. at base, jorquettes 
arising symmetrically, flowers (yellow) and fruit on trunk and main branches, 
mature fruiting peduncle 4 cm. long, 1 cm. thick, pod 26-28 x 10-11 cm., bluntly 
ridged, 23 VI 1953, Holliday 140 (COL, TRIN, US). La Elsa; tree 9-12 m., 
30 cm, in diam. at base, jorquettes arising symmetrically, flowers yellow, fruit on 
trunk and main branches, 23 VI 1953, Holliday 139 (TRIN, US)* Hoya del Rfo 
Sanquininf, left side, La Laguna, forests 1250-1400 m. alt.; tree 20 m. alt., yellow 
flowers, fruits large, brown tomentose, "cacao indio," 20 XII 1943, Cuatrecazas 
15700 (F, VALLE, Y). 
12. Theobroma stipulatum Cuatr. Figures 30, 32, 35, 37; Map 9 
Theobroma stipulatum Cuatr. Fieldiana Bot. 27(1): 84, fig.7. 1950; Baker, 
Cope, et al. (1954) 14, line 15 (as Theobroma sp.); Le6n (1960) 322, 315, 
fig- 
Type.—Ouatrecasas 21339, Colombia, Choc6. 
Large tree to about 30 m. high; growth pseudoapical; trunk about 45 
cm. in diameter, somewhat triangular at base, the bark rugose, 
granulate, reddish brown, the wood dark ochraceous, hard; branches 
grayish, rugose-squamulose, glabrate, the primary ternate; terminal 
branchlets pale ferruginous tomentose, densely covered by stellate or 
fasciculate hairs; stipules coriaceous, densely tomentose, pale fer- 
ruginous, ovate or ovate-oblong, obtuse, persistent, 8-12 mm. long, 
5-9 mm. broad, the terminal up to 25 x 11 mm. 
Leaves large, strongly coriaceous; petioles robust, very thick, 
short, densely ferruginous tomentose, 5-10 mm. long; blades ovate- 
elliptic or elliptic, more or less oblong, rounded, truncate or obtuse at 
apex, the usually slightly asymmetrical base emarginate-cordate, 
entire or slightly sinuate and flat at margin, 23-45 cm. long, 11-17 cm. 
broad, green above, when dry brown or pale brown, slightly rugose or 
542 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
almost smooth, with scattered, minute, stellate hairs, the costa and 
secondary nerves linear, tomentose, depressed, greenish ochraceous 
beneath or when dry ochraceous brown or pale brownish, tomentose, 
covered with minute, whitish, intricate, stellate hairs, and other 
mediocre, thicker, ferruginous, stellate hairs copiously covering the 
veins, the costa very robust and prominent, the secondary nerves 
about 12 on each side, very prominent, sub ascending, near the margin 
curved, decurrent, and anastomosing, the transverse tertiary nerves 
prominent, parallel, 5-15 mm. distant from each other, the minor 
veins prominously reticulate. 
Inflorescences caulinc, on trunk or main branches, the fertile 
branches perennial originating from tubercles; ligneous branches 
short, tortuous, intricate, furcate-ramose (dichasial and cincinnate), 
bracteate, tomentose, up to 1-3 cm. long; bracts ovate, sub coriaceous, 
tomentulose, minute; peduncles solitary, 5-12 mm. long, thin, stellate- 
tomentose, 3 bracteolate and 1 -flowered at apex; bracteoles linear, 
acute, 2-3 mm. long; pedicels erect, thin, densely tomentose, 10-22 
mm. long; buds globose, 10-14 mm. in diameter, sublanate-tomentose; 
sepals thick, ovate-triangular, glabrous and ochraceous inside, except 
for the minute, thick, oblong, glandular hairs at base, densely stellate- 
tomentose or sublanate outside, 10-15 mm. long, united in the lower 
third, refiexed at an thesis, umbilicate at base; petals yellow, glabrous, 
the hoods obovate elliptic, 7-nerved-sulcate, involute at margin, 
round-cucullate at apex, about 5 mm. long and 3-4 mm. broad; 
petal-laminae yellow, thick, 3-3.5 mm. long, ca. 3-3.5 mm. broad, 
suborbicular or subspatulate, slightly rctuse at apex, attenuate into a 
narrowly linear pedicel at base, this 3.5-4 mm. long. 
Androecium tube 1.5-2 mm. long, glabrous; staminodes petaloid, 
yellow, thick, glabrous obovate-oblong, subspatulate, rounded at 
apex 1-toothed on each side, 10-11 mm. long, 4.5-5 mm. broad, at 
base 1.5 mm. broad; filaments rather thick, glabrous, 1.8-2 mm. long, 
curved, shortly 3-furcate, 3-antheriferous; anther lobes ellipsoid; 
ovary ovoid-oblong, 5-furrowed, densely tomentose-hirsute, about 2 
mm. high; styles filiform, glabrous, coherent, about 2 mm. long. 
Fruit 17-22 cm. long, 9-11 cm. broad, ovoid-ellipsoid or ellipsoid 
and oblong, rounded at base, slightly attenuate, obtuse or rounded at 
apex; pericarp hard-coriaceous, rigid, smooth, appressed brown 
tomentose, the epicarp woody, about 1.5 mm. thick, the mesocarp 
and endocarp carnose, creamy, about 1 cm. thick; seeds compressed 
about 20-55, surrounded with pale, yellowish white, soft, scented 
pulp, more or less amygdaliform, 20-25 mm. long, 18-21 mm. wide, 
and 7-10 mm. thick, the testa sub coriaceous about 0.5 mm. thick; 
cotyledons white; germination hypogeous; fruiting peduncles robust, 
1-3 cm. long, about 1 cm. thick. 
CUATRECASAS—CACAO AND ITS ALLIES 543 
Common names,—Chocolate de monte, cacao de monte. 
Uses.—The seeds are said to yield a good chocolate but they are 
only occasionally used by the natives. 
Distribution.—Restricted to the rain-forested basins of the rivers 
San Juan and Atrato (Choc6 region) and perhaps Rio Sequi6n (Narifio) 
in western Colombia, where it is of rare occurrence. 
COLOMBIA: Antioquia: Villa Arteaga, about 100 m. alt.; tree 12 m., 20 cm., 
diameter base, flowers borne on trunk and main branches, pedicel from base 
to bracts 1.6 cm., from bracts to flower 2.2 cm., bracts 3, one usually larger, 
abscission layer near bracts, sepals joined about % way from base, reflexed when 
flower opens, all parts of flower yellow, 7 ridges on inside of petal, staminodes 
spatulate, 14 mm. long, 6 mm. wide, 22 VII 1953 Holliday <fc Bartley 7/165 (TRIN, 
US). Ibidem, and same tree; Cope Ant. 2 (specimens lost), field annotations 
by Cope: "A branched tree, showing the branching of the subincanum group, new 
growth from above jorquette; 3 branches." "Interior parts of bud examined 
were creamy white; sepals reflexed in opened flower, 7 mm. long x 4 mm. wide, 
elliptic, pointed, apparently rather soft and spongy, truncate margins, free almost 
to base, inner surface glabrous; petals 5, free, with cup-shaped base and ligule, 
base 4x4 mm., ligule and strap 5 mm. long, expanded portion spatula-shaped, 
staminodes 5, reflexed, oblanceolate, 10 mm, x 3 mm., fruit somewhat ovate in 
outline, in cross section, slightly flattened on 5 side, no ridges or furrows, densely 
covered with medium brown, stellate hairs, which impart a mealiness to surface, 
18 cm. long x 10 cm. diameter, wall with woody outer layer about 1.5 mm. thick, 
inner surface very soft, creamy, about 1 cm. thick, 55 seeds embedded in soft, 
cream-colored tissue, beans flattened, up to 1 cm. thick, somewhat triangular in 
outline, up to 25 mm. long x 21 mm. wide, testa chocolate brown in color, about 
0.5 mm. thick, rather leathery, cotyledons pure white, markedly convoluted, pulp 
has smell of green bananas." 
Choc<5: Rio San Juan, right margin of river on low hill near Falestina, about 
30 m. altitude; tree 32 m. tall; trunk triangular at base, 45 cm. diam., branchless 
up to 25 m. height, bark rugate-granulate reddish brown, wood rather dark 
ochraceous, leaves coriaceous, rigid, green above, ochraeeous green beneath, when 
very young light yellowish green above, young branchlets and stipules pale 
ferruginous, fruits oblong-cllipsoid or oblong-subovoid, obtuse, dark brown 
tomentose, 17-22 cm. long by 9-10 cm. broad, pericarp rigid woody, "chocolate 
de monte," 28 V 1946,Cuatrecasas 21339 (holotype, P; isotypes, US, VALLE, Y). 
Rio Atrato, Llord; young tree 3 m., sterile, jorquette of three branches, 4 VIII 
1953, Holliday & Bartley 175 (TRIN, US). 
NakiSTo: Iscuand6, Rfo Sequrtn, 100 m. alt.; tree IS m., bark reddish and 
smooth, wood cream colored, "chocolate," 23 XI 55, Romero Castaneda 5500 
(COL). (A sterile specimen, the identification is doubtful; flowers and fruits 
necessary.) 
13. Theobroma chocoense Cuatr., sp. no v. Figures 3, 31, 36, 38; Map 9 
Arbor 8-15 m., trunco erecto pseudoapicale crescente, cortice sub- 
laevi, ramis ternatis robustis patulis superioribus ascendentibus ramulia 
alternis juvenilibus dense crasseque lanuginoso-tomentosis viridi- 
ochraceis vel viridi-ferrugineis, pilis stellatis intricatis mediocribus 
ochraceis densis et pilis fasciculatis longioribus pallidis sparsis tectis, 
denique glabratis brunneis nitidis angulosis cicatricosis; stipulae 
coriaceae persistentes petiolis longiores ovatae obtusiusculae longi- 
544 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
tudinaliter striato-nervatae primum ochraceo-tomentosae, pilis fer- 
rugineis mediocribus stellatis, delude cinereae, pilis stellatis minutis 
albidis adpressis munitae, intus glabrae 12-22 mm. longae, 10-15 
mm. latae, in ramis valde juvenilibus ovato-lanceolatae 2-25 mm. 
longae, 8-9 mm. latae. 
Folia alterna rigide coriacea; petiolus brevis crassus robustus viridi- 
ockraceus vel viridi-ferrugineus dense lanuginoso-tomentosus, pilis 
mediocribus stellatis densis et pilis fasciculatis longissimis (ad 2 
mm. longis) pallidis sparsis, 7-14 mm. longus, 8-10 mm. crassus; 
lamina oblongo-elliptica vel subovato-elliptica, Integra vel Ievissime 
undulata, basi ampla leviter attenuata subsymmetrica rotundata cor- 
dato-emarginata, apicem versus paulo attenuata apice obtusa subite 
acuteque acuminata 18-42 cm, longa, 9-20 cm. lata, acumine 1-1.5 cm. 
longo, supra juventute ochraceo-stellato-tomentosa deinde glabra vel 
sparsissimis pilis stellatis mediocribus munita, plus minusve venoso- 
rugosa, nervis secundariis filiformibus impressis, nervulis venulisque 
plus minus ve impressis, superficie lutescenti-viridi in sicco brunne- 
scente, subtus valde nervosa viridi-cinerea in sicco ochraceo-ferruginea, 
costa crassa valde eminente, nervis secundariis valde prominentibus 
12 vel 13 utroque latere paulo ascendentibus ad marginem arcuatis, 
nervis tertians transversis parallelis prominentibus 3-10 mm, inter se 
distantibus nervulis minoribus venulisque bene prominentibus reti- 
culatis, indumento he tero tr ich o- to men toso, superficie alveolorum 
venulisque reticuli pilis-s tell at i s minutissimis tenuibus albidis in- 
tricatis dense tectis, nervis alteris copiosis punctis callosis rubris et 
copiosis, pilis stellatis majoribus ferrugineis radiis patulis munitis, 
in foliis vetustis nervis majoribus saepe glabratis, in prole foliis tenui- 
oribus subtus albo-cinereis, pilis minutissimis stellatis albis dense 
tecta, in nervulis sparsis in petiolo costa nervisque principalibus 
densis pilis stellatis ochraceis radiis patulis gracilibus 1-1.5 mm. 
longis basi calloso-rubescente praedita. 
Folia prolis orthotropae tenuia petiolata late ovata vel rhomboideo- 
ovata sursum crenato-dentata apice acuminata, circa 8 nervis secunda- 
riis utroque latere; lamina 12-20 cm. longa, 8.5-12 cm. lata, petiolo 
3,5-4 cm. longo, vel latiore; stipulae anguste lanceolatae. 
Inflorescentiae in trunco et in ramis majoribus copiosae, tuberculis 
lignosis persistentibus orientes, ramulis brevissimis cymosis tomentosis 
densos glomerulos floriferos formantibus; bracteae linearcs circa 5 
mm. longae ferrugineo-tomentosae; pedunculi ad 1 cm. longi tomentosi 
apice 3-bracteolati; bracteolae anguste lineares 2-2.5 mm. longae 
tomentellae; pedicelli 9-14 mm. longi erecti mediocres stellato- 
tomentosi viridi-ochracei; alabastra ellipsoideo- (globoso) -depressa 
9-10 mm. lata, 7-8 mm. longa, in commissures 5-costata, minute ochra- 
ceo-tomentosa; calyx basi cupularis, intus basi ad marginem insertionis 
CUATRECASAS—CACAO AND ITS ALLIES 545 
annulo pilis densis crassis glandulosis praeditus, in anthesin reflexus; 
sepala 5 crassuiscula ovato-acuta, 10-11 mm, longa, circa 7 mm. 
lata, basi 2.5-3 mm. longe in tubum coalita, extus minute ochraceo- 
stellato-tomentosa, margine minutissime albo-tomentosa, intus glabra 
superne parce pilosula excepta. 
Fetala glabra, cucullo crassiusculo pallido obovato-suborbiculato 
circa 6 mm. longo et lato, basi amplo apice subrotundato depresso 
apiculo bidenticulato, venulis 7 extus paulo conspicuis intus promi- 
nentibus; lamina crassa rigida rubra obovato-deltoidea apice sub- 
truncata, 5-6 mm. longa, 5-6.5 mm. lata, basi attenuata et cum 
pediculo circa 3.5 mm. longo articulata. 
Androecii tubus crassiusculus glaber circa 1.5 mm. altus; stami- 
nodia petaloidea crassiuscula glabra in anthesin erecta obovato-oblonga 
apice rotundata margine integra basim versus gradatim attenuata, 
13-15 mm. longa, 5-7 mm. lata, basi 3-3.2 mm. lata; stamina Ala- 
men tia crassiusculis circa 2 mm. longis recurvatis glabris, anther is 3, 
lobis bilocularibus loculis ellipsoideis circa 0.6 mm. longis; ovarium 
globosum 2.5 mm. diam. dense hirsuto-tomentosum; styli circa 2 
mm. longi. 
Fructus laevis ellipsoideo-ovoideus basi late rotundatus apice paulo 
attenuatus obtusus, 19-20 cm. Iongus, 10-11.5 cm. latus, pericarpio 
crasso duro, extus crasse brunneo-tomentosus, epicarpio circa 2 
mm. crasso lignoso; semina ovoidea vel triangulari-ovoidea compressa, 
20-23 mm. longa, 16-18 mm. lata, 11-12 mm. crassa. 
Type in the U.S. National Herbarium, No. 2402158, collected on the 
grounds of the Experimental Station " Agroforestal del Calima," 
Bajo Rio Calima, at about 35 m. altitude, in the Department El 
Valle, Colombia, September 25, 1961, by J. Cuatrecasas and L. 
Willard (No. 26074). Flowers from the same tree collected by H. 
Guerrero A. (No. 26074) December, 1961. Paratype collected at 
the same place with a dry mature fruit by V. M. Patino (No. 115). 
Common names.—Cacao de monte, cacao grande de raonte, bacao 
de monte. 
Distribution.—Restricted to the Chocd region, in the heavy 
forested valleys of the rivers San Juan and Atrat-o in western Colombia. 
The records of T. simiarum from the Calima River by Baker, Cope & 
al. (1954, 14) refer to this species. 
Theobroma ckocoense is closely related to T. simiarum, from which 
it differs in its broadly ovoid or ellipsoid pods, in its elliptic or ovate 
leaves (instead obovate), in the indument of the lower side of the 
leaves, which is composed of two kinds of hairs (minute, entangled, 
white, stellate hairs covering the surface and larger ones with long, 
spreading rays on the nerves), by the short, ovate, obtuse stipules, 
and shorter bracteoles. All these features indicate that Chocoan 
546 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
species is distinct from the Costa Rican species. Although the fruit 
of T. chocoense is very similar to that of T. 8tipulatumt the red color 
of the flower and the different indument of the leaves readily dis- 
tinguish these two species. 
COLOMBIA: El Valle: Baja Calima (Choc6 region), Rio Calima, Estaci6n 
Agroforestal del Calima, 30-40 m. alt.; tree about S m. high, with apical growing 
trunk almost smooth with few fertile tubercles, primary branches ternate, abun- 
dantly inflorescence-tuberculate, leaves rigid, medium yellowish green above, 
cinereous green beneath, "cacao de monte," 25 IX 1961, J. Cuatrecasas L. 
Wiltard 26074 (holotype, US; isotype, COL). Ibidem; flowers purple red, XII 
1961, Humberto Guerrero 26074 (US). Ibidem; small tree (about 10 ra.) on the 
edge of the small creek behind the Palmetum, dry fruit 19 x 11.5 cm., I 1953, 
V. M, Patino 115. Ibidem; tree 12-15 m. in forest, fruit 16 x 11 and 20 x 10 
cm., quite smooth, no flowers, 29 VI 1953, Holliday 144 (TRIN, US). Rio 
Calima, Caflo la Brea; young tree 2 m., in forest, sterile, 29 VI 1953, Holliday 143 
(TRIN, US). 
Choc6: Rio San Juan, in front of Palestina, Quebrada de la Sierpe, 0-40 m. 
alt.; tree about 14 m. high, trunk 30 cm. diam., leaves rigid, rugose, coriaceous, 
deep green above, cinereous green beneath, strongly nerved, fruit ellipsoid, 
smooth, velvety tomentose, 20 cm. long, 10 cm. wide, the seeds acid, "bacao de 
monte," 13 III 1944, J. Cuatrecasas 16896 (F, VALLE, US). 
13a. Theobroma chocoense var. bullalum Cuatr., var. nov. 
A var. chocoensi foliis rugosis reticulato-bullatis differt. Lamina 
rigid a elliptico-oblonga vel paulo obovato-oblonga, magna, usque ad 
50 x 21 cm., basi rotundata, apice rotundata breviter subiteque cuspi- 
data, juvenilis basim versus plus minusve attenuata apice magis 
angustato-acuminata; cucullum petalorum 5-6 mm. longum, 3-4 mm. 
latum, pediculo 3,5-4.5 mm. longo; lamina triangulari-spathulata, 
2.5-4 mm. longa, 2,5-3 mm. lata; staminodia obovato-oblonga apice 
rotundata, circa 10 mm, longa, 4 mm. lata, basi 1.5 mm. lata; sepala 
extus tomentosa intus glabra, circa 12 mm, longa, 5 mm. lata, 3 mm. 
basi coalita. 
Type in the U.S. National Herbarium, No. 2404636, collected in 
Quebrada Juan Maria, a small tributary of Rio Jurad6, at about 500 
m. from its mouth, in the Municipality of Nuqui, Department of 
ChoctS, Colombia, at 50-100 m. altitude, on a hill in a region having a 
dry season of about 4 months, 15 IX 1955, V. M. Patino 171. 
Although at first sight the leaves of this varietyflook very different 
from typical T. chocoense, no other differences can be found on the 
basis of the available material. According to fragmentary data it 
seems that the fruit of the Juradd specimens are identical with those 
of T. chocoense. The leaves are strongly nerved, reticulate and rugose- 
bullate, but some specimens of typical T. chocoense also show rugosity 
in the leaves. V. M. Patino writes about his specimens: "Tree 12-14 
m. alt., the trunk straight, triangular on the lower half, with sides 
CUATRECASAS—CACAO AND ITS ALLIES 547 
about 30 cm. broad, tubercular, floral cushions copious on the whole 
trunk; the specimens are scattered; no other trees were seen in the 
vicinity; the "cuzumbies" like its fruits very much, for which reason 
it is difficult to find them unbroken; branches few, terminal; leaves 
40-50 x I672I cm., petiole 1-1.5 cm., 6-11 mm. thick; leaves of very 
young specimens dentate in their upper half; fruits on trunk and 
branches; the 2 flowers collected were dry but a Choko Indian told 
that its natural color is red and that the seeds are purplish; the Choko 
name for the species is "cumajti" or "judromajd." 
To this variety belong the sterile specimens from Rio Atrato referred 
to by Baker, Cope and al. in their Report, page 14 as "Theobroma sp. 
(possibly new)." Additional flowering and fruiting collections of 
this plant are needed to complete our knowledge and to determine 
better its taxonomic rank. 
Common names.—Chocolate de monte, bacafto de monte, cumaj6 
(Choko), judromaj<5 {Choko), according to Victor M. Patino notes. 
Uses-—The pulp is acid and of pleasant taste, for which reason it is 
sought by animals and also by the Indians. 
Distribution.—Rio Atrato and northwestern Pacific drainage, 
Choc6 (Colombia). 
COLOMBIA: Choco: Rfo Atrato, Llor6, about 50 m. alt.; tree 12 m., 4 VIII 
1953 Holliday & Bartley T/176 (K, TRIN, US). Municipality Kuqul, Rfo Jurad<5, 
Quebrada Juan Maria, about 500 m. from mouth, 50-100 m. alt., 15 II 1955, 
Patino 171 (US). Ibidem, seedling, Patino 171A (US); first branches of a 
young plant, Patino 171B (US). 
14. Theobroma siimarum Donn. Smith. Figures 5, 6, 32, 36, 38, 40; Map 8 
Theobroma simiarum Donn. Smith in Pittier, Prim. Fl. Costar. 2: 52. 1898; 
Bot. Gaz, 25: 145. 1898; DeWildeman (1902) 97; Standley (1937) 689; 
Chevalier (1946) 282; Holdridge (1950a) 2; Allen (1956) 343; Le6n (1960) 
322, 321,fig. 
Type.—Turrialba, Costa Rica, Tonduz 8373 (= 7313 distributed 
J. Donnell Smith). Lectotype: US 1,382,332 [Photo F. M. 40723], 
US 471,873 [Photo F. M. 40722]. Syntypes: Pittier tfb Tonduz 
3925, Tonduz 6852, Cooper 10244. 
Medium to large tree up to 20 m. tall, with erect, thick trunk 
up to 60 cm. in diameter at the triangular base; growth pseudoapical; 
bark more or less rugose; branches spreading, the primary ternate, 
the upper ones ascending; young branchlets thick-tomentose, greenish 
ferruginous or brownish, covered by three kinds of hairs: 1) abundant 
minute squamose stellate hairs with thin rays, 2) mediocre stellate, 
longer hairs and 3) scattered large stellate or furcate hairs with very 
long white rays, when older glabrate, brownish or grayish; stipules 
persistent, longer than the petioles, subcoriaceous, firm, erect, lanceo- 
548 CONTRIBUTION'S FROM THE NATIONAL HERBARIUM 
late or linear, acute, 15-24 mm. long, 3-6 mm. broad, tomentose 
outside. 
Leaves large, rigid, coriaceous; petiole robust, thick, somewhat 
striate, thick-tomentose-ferruginous, 5-12 mm. long, 6-10 mm, thick; 
blades obovate-oblong, rounded or very obtuse and abruptly cuspidate 
at apex, broad or slightly narrowed, rounded or slightly cordate and 
asymmetrical at base, entire or sinuate-dentate in the upper third, 
in very young plants obovate-rhomboid, long-acuminate, the upper 
half or third repand, acutely dentate, the adult 20-40 cm. long, 8-17 
cm. broad, the acumen 0-1 cm. long, green above, when dry pale 
brown, smooth, glabrous or with scattered stellate hairs on the midrib, 
this and the secondary nerves filiform and depressed, the other veins 
usually obsolete, light green beneath, when dry cinereous or whitish 
but the nervation ferruginous, the costa thick, very prominent, the 
9-11 secondary nerves on each side thick and prominent, spreading- 
ascending, the lower pair usually at a more acute angle, the transverse 
tertiary nerves prominent, separated 5-10 mm. from each other, 
the minor nerves and veins prominulous, reticulate; indument hetero- 
trichous, the areolar surface and the small reticular veins covered by 
a dense tomentum of intricate, minute, white, stellate hairs, the 
thicker nerves with sparse mediocre, ferruginous, stellate hairs and 
reddish callous scars, the costa densely ferruginous by more or less 
copious mediocre hairs and other ochraceous, long ones; in seedlings 
the under leaf surface whitish, covered with a white tomentum of 
extremely minute, white, stellate hairs, and on the nerves sparse an d 
on the costa dense ferruginous stellate hairs with long (1 mm.), thin, 
patulous rays and red, callous bases. 
Inflorescences borne on tubercular protuberances on the trunk; 
cymose branches usually cincinnate, ligneous, very short, forming 
dense, many-flowered glomerules; branchlets up to 4 cm. long, tor- 
tuous, rugose, glabrate, brae tea te, the bracts coriaceous, ovate, 
subacute or acuminate, tomentulose outside, 1-2 mm. long, 1-1.5 mm. 
broad, persistent; hornotinous branchlets short, crowded, ferruginous- 
tomentose, covered by fertile, tomentose, imbricate bracts; pe- 
duncles axillary, mediocre, densely stellate-pilose-tomentose, 5-15 
mm. long, 3-bracteolate at apex, the bracteoles narrow-linear, to- 
mentose outside, 3-7 mm. long; pedicels up to 15 mm. long; buds 
globose, densely tomentose, above almost 5-angulate; calyx cupular 
at center, subglabrous, with a ring of minute, thick, oblong, glandular 
hairs at base and with very thin hairs above inside, densely and 
thickly stellate-tomentose and ochraceous green or ferruginous outside, 
the sepals ovate-acute, about 10-12 mm. long, 6 mm. broad, united 
about 2 mm. at base, reflexed at an thesis; petals red, thick, glabrous; 
hood yellowish white, obovate-oblong, with 7 prominent nerves inside, 
CUATRECASAS—CACAO AND ITS ALLIES 549 
v 
Figure 38.—Leaves of Theobroma: a, simiarum, from orthotropic stem (Stand. 37377); 
b, simiarum, from normal lateral branches (Cooper 10244); c, chocoense (Cuatr. 26074); 
d, leaf base and stipules of T. chocoense (Cuatr. 26074). All leaves X d, X %• 
550 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
narrowed-unguiculate, and inside reddish at base, 6-8 mm. long, 4-5 
mm. wide, rounded-cucullate, inflexed-apiculate at apex, the apiculum 
bifid, articulate to the pedicellate lamina; petal-lamina obtrapezoid, 
carnose, red, subtruncate at apex, cuneate at base, 3.5—4.6 mm. long, 
3.5-5 mm. wide, refiexed in bud, erect in an thesis; pedicel linear, 
4-5 mm. long, 1 mm. broad; androecium tube about 2 mm. long, 
rather thick, glabrous; staminodes petaloid red, glabrous, obovate- 
oblong, rounded at apex, slightly sinuate-dentate in the upper third 
or subentire, 11-13 mm. long, 5-7 mm. broad, 1-2 mm. wide at base, 
reflexed in bud, erect at anthesis; filaments rather thick, glabrous, 
curved, 2 mm. long, shortly 3-furcate, 3-antheriferous; anther lobes 
ellipsoid, 0.6-0.7 mm. long; ovary obovate, densely hirsute-tomentose, 
about 1.2 mm. high; styles about 1.2 mm. long, thin, acute, united 
at base. 
Fruiting peduncle short, thick, usually 1-1.5 cm. long; fruit ellip- 
soid-oblong, smooth, truncate or very blunt, rarely narrowed at 
base, rounded, very obtuse or slightly attenuate at apex, 16-35 cm. 
long, 6-10 cm. broad, covered by dense and thick, ferruginous or 
brown tomen turn; pericarp 11-15 mm. thick with: 1) epicarp woody, 
1.5-2 mm. thick, with a tomentose epidermis, 2) mesocarp 5-10 mm. 
thick, firmly carnose, pale ochraceous whitish, and 3) endocarp 2-5 
mm, thick, softer carnose; seeds 36-60, distributed in 5 more or less 
double rows, each one covered by a thick, fibrose, white or rosy- 
white, aromatic pulp irregularly ovoid, 20-21 mm. long, 18-17 mm. 
broad, and 13-15 mm. thick; cotyledons white; germination hypogeous. 
Common names.—Cacao de mico, cacao de mono, teta negra. 
Indian names: Kr&aku (Guaiuso), Nun (sup (llama), Uirub (Bribri), 
Dzug-mang-ua (Brunka), Ku-gin, Bik (Terraba), according to 
Standley, loc. cit., Uir-ub (Bribri) according Pittier. 
Uses.—The seeds are said to give a cacao of good quality. They 
are occasionally used. 
Distribution.—Limited to Costa Rica, Infrequently found in 
preserved or remnant forests in both Atlantic and Pacific lowlands, 
from sea level to 600 m,, and exceptionally in higher altitudes up to 
900 m.; recorded from the provinces of Lim6n, Heredia, Puntarenas, 
and Cart ago. It may extend to Nicaragua and northern Panama, 
but no records from these two countries exist at present in herbaria- 
In its natural habitat T. simiarum attains a considerable height 
and thickness. The trunk is erect and branchless up to several meters. 
The inflorescences may appear on the trunk but they are more abun- 
dant on the big branches. The fruit is very characteristic because of 
its terete, sausage shape. From the closely related Colombian species 
T. stipulatum and T. chocoense it can be distinguished by the elongate 
CtJATRECASAS—CACAO AND ITS ALLIES 551 
shape of the pod; in the two other species the fruit is ovoid or broadly 
ellipsoid. From T. stipulatum it differs also by its red flowers, longer 
lanceolate stipules, longer bracteoles, and by the indument of the 
leaves. From T. ckocoense it differs also by its narrower and longer 
stipules, the obovate form of the leaves, by the leaves having beneath 
usually three kinds of hairs, and by its longer bracteoles. 
COSTA RICA: Cartago: Turrialba, grassland, 570-600 m. alt., XI 1893, 
Tonduz 8373 (distributed by Donn. Smith under number 7313) Iectotype US; 
isolectotypes BR, G, OH, K; Photos F. M. 30722, 40723, 40743. Turrialba, forest 
at margin of Rfo Reventaz<5n, about 600 m. alt.; tall tree with unbranched trunk 
in the lower half, 60 cm, diam. near the subtriangular base, abundant tuberculate 
inflorescences on the upper part and big ternate branches, few sausage-shaped 
pods, 6 XI1961, J. Cuatrecasas <fc Ledn 26515A (US), Turrialba, Institute Inter- 
americano C.A., 600 m. alt.; flowers bright red, cult., 4 XI 61, J. Cuatrecasas & J. 
Le6n 26515 (US). Turrialba, I.I.C.A., 600 m. alt., cult., 14 III 51, J. Le6n 
3189 (TURRI). Tuis, forests, 670 m. alt., "cacao de mico," XI 1900, Pittier 
14016 (Donn. Smith 7731) (GH, K, US). Tucurrique, grassland and forests 
around Las Yueltas, about 635 m. alt.; large tree, fruits brown, oval-elongated 
30-40 cm. long, XI 1898, Tonduz 12822 (BM, G, LE, P, US), 18222 (M) [erroneous 
number]. 
Limon: La Colombiana Farm of the United Fruit Company, about 70 m., 
wet forest; tree 60 feet or more tall with trunk 2 ft. thick, small crown, flowers 
bright red in bunches on trunk, fruit sausage-shaped, hairy, 1 foot long or more, 
said to be rare here, seeds give good cacao, 6, 7 III 1924, Standley 36822 (US). 28 
miles on the railroad from Puerto Lim6n, towards Rio Barbilla, in marginal 
forest, about 60 m. alt.; tall tree, flowers flame red, 12 V 1930, Cufodontis 599 
(WU). Along Rfo Reventazdn, below farmhouse Finca Castilla, 30 m. alt., in 
Gynerium sagittatum thickets, 27 VII 1936, Dodge & Goerger 9420 (MO). Palm 
swamp between Rfo Reventaz6n and Rfo Parismina, on Castilla Farm, 2 IV 
1930, Dodge & Nevermann 7164 (MO). La Lola, farm of the I.I.C.A., about 40 
m. alt., cult.; trees about 10 m. high, trunk 20-30 cm. diam., triangular at base, 
flowers scarlet, fruits oblong 15.5 x 7 cm., 18 x 8.7, 19.5 x 8.5, 22.7 x 7.7, 23.3 x 
8.4, 25.7 x 9.3 cm., with thick brown tomeutum, 6 XI1961, Cuatrecasaa & Paredes 
26536 (US). Vicinity of Gudpiles, 300-500 m., seedling, III 1924, Standley 37377 
(US). 
Heredia: La Concepci6n, Llanuras de Santa Clara, 250 m. alt., "cacao de 
mico," II 1896, Donn, Smith 6457 (BM, US). Santa Clara; 6-7 m. high tree, 
"cacao de mono," IX 1896, Cooper 10244 (syntype, US) (Photo F. M. 40724). 
Puntarenas: Terraba, 260 m. alt., forests; caulinar flowers, large cylindrical 
fruits (30-35 by 10 cm.), "cacao de mico," II 1891, Pittier & Tonduz 3925 
(syntype, BR). Boruca, forests 466 m. alt., Ill 1892, Tonduz 6852 (syntype, US). 
TRINIDAD (cult.): Imperial College of Tropical Agriculture, Cuatrecasaa & 
Cope 25792 (US), 25794 (US). 
SURINAM (cult.): Paramaribo, culture garden; tree 10 m., 15 cm. diam., 
almost horizontal branches, flowers bursting in dense clusters from the trunk 
and old branches, calyx rusty brown, ligula and staminodes shining lacquer red, 
8 IV 1954, Idndeman 5725 (U). Ibidem; seeds round, cotyledons white, germina- 
tion hypogaeic, VIII 1956, van Sucktelen, s.n. (US). 
BRASIL (cult.): Parfi, cultivated and said to be introduced from Venezuela; 
small tree with purplish flowers, II V 1957, Pires 6575 (IAN). 
680-695—64 12 
552 
15. Tbeobroma grandiflorum (Willd. ex Spreng.) Schum. 
Figures 3, 5, 6, 7, 33, 36, 39, 40; MAP 10; PLATE 8 
Tkeobroma grandiflorum (Willd. ex Spreng.) Schum. in Mart. Fl. Bras. 
12(3) :76, pi. 8, 1886; Jumelle (1899) 28, Jigs. 14, IS; De Wilde man 
(1902) 95; Ducke (1925) 131; Ducke (1940) 272, pi 4, fig- Chevalier 
(1946) 281; Addison & Tavares (1951) 25, pi. l,fig. 1, pi. 8, fig. A, pi. 14* 
jig. 10; Ducke (1953) 11; Baker, Cope & al. (1954) 13, fig. 14; Cuatreeasas 
(1956) 656; Le6n (1960) 320, 319, fig. 
Bubroma grandiflorum Willd. ex Spreng. Syst. Veg. 3:332. 1826. 
Quazuma grandiflora (Spreng.) Don, Hist. Dichl. 1:523. 1831. 
Theobroma speciosum Willd.?, sensu Mart, in Buchn. Repert. Pharm. 35:24. 
1830; Linnaea Litt. Bericht 32. 1831, non Willd. 
Tkeobroma macrantha Bernoulli, Uebers, Art. Theobroma 11. 1869. 
Theobroma silvestre Spruce ex Schum. in Mart. Fl. Bras. 12(3): 76. 1886, 
as synonym. 
Type.—Siler 4, "Hoffmannseg" in Herbarium Willdenow No. 
14352 (B). Spruce 1822 (syntype of T. macrantha Bernoulli and 
neotype of B. grandijlorum Willd.). 
Medium-sized tree, usually 6-10 m. high, reaching up to 18 m.; 
growth pseudoapical; trunk up to 25-30 cm. in diameter, the bark 
grayish, granulose, more or less wrinkled, internally rosy or reddish, 
the wood pale; branches robust, spreading, the superior ascending, the 
primary ternate; branchlets terete, the terminal rather thick, densely 
and abundantly lanate-tomentose, the indument ferruginous, floccose, 
Map 10.—Geographical distribution of Theobroma obovatum # and T. grandiflorum Q< 
CUATRECASAS—CACAO AND ITS ALLIES 553 
Figure 39.—Leaves of Tkeobroma, X A> obovatum (Poeppig 8,n.); b, grandiflorum (Cuatr. 
25780T); c, grandiflorum (Killip & Smith 30011); d, kylaeum (Araque & Barkley 18C745); 
E, subincanum (Baker 38). 
554 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
more or less deciduous, the hairs stellate or fasciculate with long, 
intricate rays, caducous, the surface fulvous, with exfoliating rhyti- 
dome, becoming gray, glabrous, and reticulate-rimose; stipules sub- 
coriaceous, rigid, oblong or lanceolate-oblong, obtuse or subacute, 
nervate-striate, abundantly lanate-tomentose, the tomentum more or 
less deciduous, 10-20 mm. long, 3-6 mm. broad, persistent. 
Leaves firmly coriaceous, medium sized or large; petiole thick, terete, 
with a dense, thick, ferruginous, lanate tomentum, 7-14 mm. long, 
2-5 mm. thick; blades oblong, subobovate-oblong or sub elliptic- 
oblong, more or less attenuate toward the base, this obtuse, rounded, 
usually emarginate, or subcordate and slightly irregular, abruptly 
attenuate and acutely acuminate at apex, the margin entire or slightly 
dentate-sinuate toward the apex, 20-35 (15-60) cm. long, 6-11 (5-16) 
cm. broad, the acumen 1-2.5 cm. long, glabrous above, green, more 
or less shining, brownish olivaceous or tabacine when dry, the midrib 
and secondary nerves filiform, depressed, the tertiary ones slightly 
marked or obsolete, greenish cinereous, glaucescent, or pale rosy 
beneath, the costa very prominent, the 9 or 10 pairs of parallel second- 
ary nerves prominent, subascending, near the margin thinner, curved, 
anastomosing, the lowest pair usually forming a more acute angle 
and more distally separated from the next, the filiform tertiary nerves 
prominent, transverse, parallel, the minor nerves and veins forming 
a fine prominulous reticulum, the costa, secondary, and tertiary nerves 
glabrous, nitidous, with sparse reddish, callose dots, the reticulum 
and areoles minutely tomentose, covered by white, minute, intricate, 
dense, stellate hairs. 
Inflorescences small, axillary and extra-axillary on leafy branches, 
the short cymes reduced to 3-5 flowers or fewer, the branchlets 
extremely short, ferruginous tomentose, the peduncles robust, 2—5 
mm. long, 3-bracteolate at apex, the bracteoles narrowly linear, 
tomentose, 3-4 mm. long; pedicels robust, rather thick, tomentose, 
ebracteate, 5-20 mm. long. 
Calyx subcymbiform; sepals firm, thick, carnose, ovate-oblong, 
subacute, about 14-15 mm. long, 6-8 mm, broad, 1.5 mm, thick, 
united in the lower third, but often the five separated to near the base 
in two pairs and one free (2S+2S-fS), the margin involute, the apex 
minutely indexed, thickly stellate-tomentose outside, ochraceous 
green or ferruginous, rosy or reddish inside, shining, minutely, 
sparsely, whitish pubescent, at base with minute, thick glandular 
trichomes, the margin densely and minutely whitish tomentose. 
Petal-hoods thick, carnose, whitish or yellowish, often with red 
lines, obovate, rounded-cucullate at apex, 7-nerved, rugulose without, 
pubescent, glabrous within, near the base reddish, 6-7 mm. long, 4-6 
mm. broad; petal-lamina dark red or crimson, pedicellate, thick, 
CUATRECASAS—CACAO AND ITS ALLIES 555 
carnose-coriaceous, trapezoid-elliptical, more or less truncate or slightly 
retuse, obcordate, minutely puberulous, 4-9 mm. long, 4.5-8.5 mm. 
wide, abruptly contracted at base into a 4.5-7 mm. long, 1.5 mm. 
broad pedicel, 
Androecium tube 2.5 mm. high, sparsely pilose; staminodes reflexed 
in bud, spreading in an thesis, crimson, dark red or purplish red, 
lanceolate, very acute, thick, but somewhat flattened with a marked 
midrib, 9-15 mm. long, 2-2.5 mm. broad throughout, pilose, especially 
outside, the abundant hairs rather long, thin, flexuous; filaments 
1.7-2 mm. long, thick, pilosulous, very shortly 3-furcate, 3-anther- 
iferous; lobes of the anthers broadly elliptic, 1 mm. long; ovary 
pentagonous-obovate, densely whitish hirsute-tomentose; styles 2 
mm. long, connivent, free to the base. 
Fruits falling from tree at maturity, without peduncle, densely 
covered with a brown tomentum, large, smooth, ellipsoid or obovoid- 
ellipsoid, rounded at both ends or, rarely, slightly attenuate at apex, 
umbilicate and conically excavate at base, 16x10-25x12 cm.; pericarp 
about 1 cm. thick, with: 1) epicarp hard, woody, about 2 mm. thick, 
covered with a thin, tomentose epidermis, 2) mesoendocarp about 7 
mm. thick, softly carnose at maturity with a thin, but firm, inner 
pellicle limiting the seed cavity; seeds about 50, 5-seriate, each 
surrounded by a yellowish, acidulous and distinctively scented, fibrous 
pulp, the inner tegument a delicate pellicle, the testa subcoriaceous, 
light brown, striped from the remains of the pulp, ovoid, or ellipsoid - 
ovoid, more or less flattened, 20-30 mm. long, 20-25 mm. broad, 
10-12 mm. thick; embryo white marbled, 19-23 mm. long, 16-20 mm. 
broad, 9-12 mm. thick; cotyledons white; germination hypogeous. 
The type of Bubroma grandiflorum Willd. ex Sprengel is the specimen 
No. 14352 of the Willdenow Herbarium in Berlin, received from the 
Hoffmannsegg Herbarium, collected in Brazil by Siber. This species 
was transferred to Guazuma by G. Don who probably did not see the 
plant, his description being taken from Sprengel. Schumann found 
out that B. grandiflorum is synonymous with T. macranthum described 
by Bernoulli many years later. The short diagnosis given by Sprengel 
agrees with T. macranihum and the description of the leaves ("amplis 
oblongis abrupte acuminatis integerrimis") disagrees with any known 
species of Quazwma, which always have serrate leaves. Freitag, who 
listed Bubroma grandiflorum as synonym of Guazuma ulmifolia 
(p. 216), did not see the type. In 1952, Dr. Mildbraed of the Berlin 
Herbarium wrote me about this type specimen: "Im Herbar Willdenow 
fehlte unerklarlicherweise der Bogen 14352 mit Bubroma grandiflorum." 
Schumann had seen the type in the Willdenow Herbarium, for which 
reason I accept his judgment as final regarding the synonymy 
established by him. Accordingly, I propose Spruce 1822 as a neotype 
556 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
of B. grandijlorum Willd. ex Spreng., as it is the first collection cited 
by Schumann and likewise a syntype of T. macranthum Bernoulli. 
Common names.—Cup ass tl, with some variations in the spelling or 
pronunciation throughout its area of cultivation, cupuagd, cupti-assli, 
copuassli, cupai-a$ti. Ducke quoted cupti do matto for wild speci- 
mens in the middle Tapajoz River, The Anglo Colombian Cocoa Ex- 
pedition recorded the following indigenous names (Baker, 1952): 
Win-che£k-ch6o-ai (Puinave), Infrida-Guaviare; bawk-pom (Maku), 
Piraparand, Tar-aira; maga (Barasana), Upper Pirap&rand; nee-aw 
(Tanimvka), Guacaya; cupu-uassti (Brazil-Portug.); ba-dja-na-hoo 
(Makuna), Lower Piraparand. 
Uses.—The natives like to eat the acid and agreeably scented pulp 
which covers the seeds, for which reason cupuassti is very much culti- 
vated or planted in the state of Para and the eastern section of Ama- 
zonas. This pulp is used to prepare soft drinks (vinho do cupuassti) 
and different kinds of preserves and candy which are exported from 
Par& and Maranhao. The taste of the pulp is sui generis, 
L. Williams compares it with that of guandbana. Patino found the 
odor of it similar to that of the "mate" (Cresceviia cujete) when be- 
ginning its fermentation. The fruits are very much liked by animals, 
especially by monkeys (macacos), which very often empty the pods 
to sip the pulp, contributing to the dissemination of the seeds. 
Distribution.—The known natural area of T. grandijlorum is 
the southern half of state of Pard, Brazil, and adjacent Amazonian 
Maranhao. It has been found wild only in the rain-forest, on ele- 
vated ground in the middle Tap aj River region (waterfalls of Man- 
gabal and of Itapacurd, Ducke), Tocantins River (railroad of Alcobaca, 
Ducke), Guamd River between Ourem and Braganga (Huber), Xingti 
River between Victoria and Alt amir a {Ducke) r and Anapti River 
(LeCdinte). The tree is always scarce in its natural area. 
This medium-sized tree with large leaves, the largest flowers in the 
genus, and the largest pods among the Brazilian cacaos, is frequently 
planted or cultivated throughout the states of Pard, Maranhao, and 
the eastern part of Amazon as to Manaos. It is also occasionally 
planted outside Brazil in warm lowlands of other tropical American 
countries, such as Colombia, Venezuela, Ecuador, and Costa Rica. 
It is found also in tropical botanical and agricultural gardens. 
MARTINIQUE: From seeds from Cayenne, L. C. Richard, s.n. (P). 
BRITISH GUIANA: "Herbarium Benthamianum," Sckomburgk, s.n. (K). 
VENEZUELA: Amazonas: Capihuara, Alto Casiquiare, 120 m. alt.; culti- 
vated, small tree up to 8-10 m., subrounded crown, trunk 25 cm. diam., gray 
bark, inner rose or reddish, wood pale, pulp of fruit used to prepare beverages 
with similar taste to the guan&bana, "cupuaau/' 28 V 1942, LI. Williams 15615 
(F, US, VEN). 
CUATRECASAS—CACAO AND ITS ALLIES 557 
COLOMBIA: Vattp^s: Monfort; tree 6 m., sepals fleshy, petals maroon, 
23 IX 1943, P. Allen 3105 (COL, MO, US). Rio Vaup&j opposite confluence 
with Rfo Papurf, Yavarat^, Salesian Mission Sao Miguel; tree about 3 years old, 
growing in full sunlight, beginning to flower, no fruit, 20 II 1952, Bartley & 
Holliday T—46 (COL, US). Rio Piraparand, near confluence with Rfo Apaporis, 
river level; young tree 3-4 m., cultivated in Indian garden, 24 VIII 1952, Baker 
& Cope 5 (TRIN). 
Amazonas: Rfo CaquetA, La Pedrera, river level; cultivated tree, rather 
exposed, in the garden of the Orfanatorio, 19 IX 1952, R. E. D. Baker 16 (COL, F, 
TRIN, US). La Pedrera, cultivated, highland; bushy tree 12 ft. tall, petals 
purple red, calyx light golden brown, staminodes yellowish, 7 X 1952, Schultes & 
Cabrera 17781 (US). Rfo Ricapuya, tributary of Rfo Apaporis, river level; 2-3 
years, young cultivated tree in Indian garden, 25 VIII 1952, Baker & Cope 6 
(COL, F, TRIN, US). Leticia, 22 VIII 1946, Black & SchuUes 46-61 (AMES, 
F, IAN, NY, U, VEN); tree 8 m., "cupu-assu/' cult., 24 IX 1946, Black & 
SchuUes 46-111 (IAN). Trapecio Amaz6nico, Amazon River, Leticia, 100 m. 
alt.; cultivated, "cupuassd," IX 1946, Schultes 8178 (AMES, F, US). 
BRAZIL: "Catal. Geogr. PI. Bras. Trop.," Burchell 9467 (GH, K, P). Ibidem, 
Burchell 9375 (syntype of Theobroma macrantha Bernoulli, K). "Amazon region," 
H. A. Wickham, s.n. (K). "Brazil Cameta," Herb. Hanbury 9471 (K). 
Amazonas: Rio Negro, Sfio Gabriel; arvore 8 m., planta antiga dos sitios, 
"cupu-assu," 27 XII1945, Fries 21556 (IAN, K, NY, USDA), Upper Rio Negro 
basin, Mouth of Rio Xie, cultivated; small tree, staminodes and ligules deep red, 
rest of flower pink, "cupu-uassu," 29 XI-7 XII 1947, Schultes & Ldpet 9204 
(AMES, F, IAN, US). Rio Negro, prope Barra; "shrub 12-15 ft., flowers only 
on ramuli, solitary normally ascendant, calyx pinkish within, petals crimson, 
cucullate, bases yellow white, coronal scales red"; the Munich specimen bears 
the number 83 on a mounting tape (F. M. Photo 40705), Oct. 1851, Spruce 1822 
(neotype of B. grandiflorum, M; isosyntypes of T. macrantha Bernoulli, BM, 
E, G, GH, K, LE, LD, NY, OXF, P, WU). Manaos, Horto Experimental, 20 
m., "eupai-ajd," 19 XII 1923, Luetzelburg 22007 (M, NY, WU). Manaos, Agri- 
cultural Experiment Station, 25 m. alt., cultivated; tree 35-40 ft., inflorescences 
on main trunk, sepals green without, pink within, petals red, 13 X 1929, Killip <fc 
Smith 30011 (NY, US). Manaos, Estrada do Aleixo, firm land; tree 6 m., flower 
red, edible fruit, "cupuagd," 16 IX 1955, Francisco (INPA) 1966 (IAN). 
Manaos, Hacienda Brasil, 15 m. alt., "cupu-affti," Luetzelburg 23287 (M). Ma* 
naos, VIII 1906, "cupn-assd," Labroy, (P). Three days upstream from Manaos, 
300 ft. alt.; grown in semishade; flowers cream colored or dark crimson, grow- 
ing out of the bark, faintly scented, Sandeman 2333 (K). Par an 6 de Matitins, 
Rio Putumayo, Iga, between TarapacA and its mouth (Santo Antonio do Iga, 
100 m. alt.); small treelet; calyx very fleshy, green yellow; staminodes white, 
laciniae flesh red turning brown on fertilization, "cupuassu," 11-18 IX 1946, 
SchuUes & Black 8146 (F, IAN). Municipality of Mahacapuru, Solinos River 
region, terra Arm a, Lago do Italiano; tree 25 ft. high, 4 inches diam., "cupu- 
assd," Krukoff 1274 (A, BM, G, K, MICH, MO, NY, P, S). Tef<5; tree 7 m., 
flower brownish rose, cult. "cupuagO," Black 47-1502 (IAN). 
Maranhao: Turyassu, Igapo-wald; Baum 6—15m., Krone gelblich, filaments 
rot-violett, 31 X 1923, Snethlage 300 (F, GH, US). 
Acre (Territorio Federal): Brasilea, in a house in front of the Bolivian town 
Cobijia, 150 m. alt., "copuassti," cultivated, 5 IX 1954, Paiino 163 (GH). 
ParA: BeWm, northeast woods of the Institute Agrondmico do Norte; 
infected with witches broom, "cupu-assu," 30 X 1942, Archer 7734 (IAN, NY, 
US). BeMm, Botanic Garden of the Museu Goeldi; native of the lower Amazon 
558 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
country; the tree becomes much larger than the cacao and the top is relatively 
narrower; the fruit resembles the pod of cacao but is much larger and the pulp 
surrounding the seeds is most delicious; Dr. Huber speaks of it as the most im- 
portant native fruit of Pard; "cupuacti," 25 VI 1908, C. F. Baker 62 (A, C, E, 
GH, LE, MICH, MO, P, U, US, NY). Museu Goeldi, 23 XI 1945, Pires & Black 
744 (IAN). Jardim Botdnico do Museu Goeldi; large tree, calyx with red marks 
inside base, spatula to portion petal dark red, fruit edible, <(cupua§ti," 11 VIII 
1942, Archer 7549 (F, IAN, K, US). Ibidem; small tree, leaves gray both sur- 
faces, flowers large, cauliflorous, calyx white yellow, staminodes deep red, petals 
pinkish, very showy, a profusion of flowers, cultivated, 15 VII 1946, Schvltes 
8065 (AMES, US). Horto Bot&nico do ParA, XI 1903, "cupuassii," Huber 4008 
(BM, G). Ibidem, "cupuassu," XI 1903, Stqueiros 4008 (MG). Bel£m, "in 
urbe cultum; arbor parva floribus brunnescentibus centro albido," "cupuassu," 
16 X 1940, Ducke 598 (F, IAN, MG, MO, NY, US). Ibidem, III V 1929, DahU 
gren & SeUa 438, 634 (F), "cupu-assu," Dahlgren & Sella 733, 739 (F, GH, US). 
Vicinity of Pari, 1 VII 1908, C. F. Baker 421 (BM, C, F, L, U, WU). In sylva 
prope Barba et alibi, VIII 1828, Riedel 1373 (A, LE, S, US). "Brasilia, Borbar," 
Riedel, s.n. (OXF). B6a Vista, on the Tapajds River, Aramanahy River, 
"cupuassti," Monteiro de Costa 121 (F). Taperinha bei Santarem, kultiviert; 
aus dem Fructen wird Marmelade bereitet, kleiner Baum, "Cupu-assti," 18 IV 
1927, Qinzberger Zerner 800 (F, WU). "Habitat in sylvis udis umbrosis ad 
Para, Dr. Martius Iter Brasil. Jul. 323," Martins [874] (M). "Prov. Paraensis 
ad Para, Dr. Martius Iter Brasil. 323," Martius [875], [876] (M) (Photo F. M. 
40706). Ibidem, Martius [873] (M) mixed with T. guianense in Photo F. M. 
19641). Par A, "Bresil- Martius," Martius (G, P). Tapana, near Par 6, woods; 
30-40 ft. tall, appendages purplish red, clearing, fruit edible, 29 X 1929, Killip 
<fc Smith 30320 (NY, US). For At des collines du Mangabal, moyen Tapaj6s, 
Cachoeira do Mangabal, "cupd do matto," 5 IX 1916, Ducke 16458 (BM, G, 
MG, P). 
Rio de Janeiro: Rio Janeiro, cultivated, "cupti assti," Glaziou 9643 (C, P). 
PERU: Loreto: Caballo Cocha, on the Amazon River, "cupuassti," VIII 
1929, LI. Williams 2401 (F). 
TRINIDAD: St. Augustine, Imperial College of Tropical Agriculture, River 
State Diego Martinez, Field 2; trunk 20 cm. diam. at base, leaves coriaceous 
dark green above, green beneath, fruits ellipsoid, densely tomentose, brown 
ferruginous, 11 x 17,11.5 x 18.5, 11.5 x 16,5, 12 x 19, 12 x 19.5, 12.5 x 19 cm., with 
47-50 seeds, pulp yellowish with special scent, cultivated, brought from Bel6m 
do Pari, 31 VIII 1961, Cuatrecasas, Cope, & Barlley 25780 T (US). Ibidem; tree 
with larger and slightly attenuate fruits at apex, 10.7 x 22, 11.1 x 21.5, 11.8 x 25 
cm., 31 VIII 1961, Cuatrecasas, Cope, & Bartley 25781 T (US). Saman Plot, culti- 
vated from seeds brought from Pard; trunk 30 cm. diam., the diameter reducing 
progressively upwards from one cluster of branches to the next, primary branching 
ternate, from the base, apical growth, bark greenish brown, granulate-lenticellate, 
minutely rimose, sepals 1.5-2 mm. thick, greenish ochraceous outside, whitish 
and pink at base inside, hoods white yellowish or dirty whitish, the margin and 
the base inside purplish, lamina car nose, dark red, later red brown, staminodes 
star-patulous, red purplish becoming red brownish, styles united whitish, ovary 
white, hirsute-tomentose, 2 IX 1961, Cuatrecasas & Cope 25801 (US). 
CUATRECASAS—CACAO AND ITS ALLIES 559 
16. Theobroma obovatum Klotzsch ex Bernoulli 
Figures 33, 35, 39, 40; Map 10 
Theobroma obovatum Klotsch ex Bernoulli, Uebers. Art. Theobroma 14, pi. 
7, Jig. 3. 1869; Ducke (1935) 132; (1940) 271, pi 5, fig. t; Chevalier (1946) 
280; Addison & Tavares (1951) 25, pi. 1, fig. 8, pi. B, fig. B, pi. 3, fig. 8, 
pi. 4tfig- Bi pi- Htfiff- 1j' Ducke (1953) 10; Cuatrecasas (1956) 657; Baker, 
Cope & al. (1954) 12, fig. 15; Le6n (1960) 322, 319, fig. 
Theobroma sylveslre sensu Huber, Bull. Herb. Boiss. II, 6: 273. 1906, non 
Mart. 
Typjb.—Maynas, Peru, Poeppig. 
Rather small tree up to 15 m. high; growth pseudoapical; trunk 10- 
25 cm. in diameter; primary branches tern ate, the inferior horizontal, 
the upper ascending; terminal branchlets slender, when young 
ochraceo-ferruginous, densely lanate-tomentose, the intricate, long, 
stellate hairs more or less floccose, deciduous, when older glabrate, 
pale brown or pale gray, the rhitidome somewhat scaly; stipules 
narrow-linear, subulate, acute, tomentose, 3-5 mm. long, 0.5-1 mm. 
wide, soon deciduous. 
Leaves chart aceous, flexible, variable in size; petiole mediocre, 
subterete, transversely rimose, densely and thickly lanate-tomentose, 
ferruginous, when old the indument appressed, grayish, 4-15 mm. long; 
blades obovate-elliptic or obovate-oblong, more or less narrowed to the 
very asymmetrical and rounded base, more abruptly attenuate and 
cuspidate at apex, entire or slightly sinuate at margin, 7 x 3 to 38 x 13 
cm., including the 0.5-3 cm. long acumen, varying very much in size 
on the same branch, when very young stellate-pilose above, then 
glabrous, pale olivaceous or pale brownish, shining, the main nerves 
depressed, filiform, the others obsolete, pale ochraceous or pale cinere- 
ous beneath, glaucescent, the costa very prominent, the 5-7 secondary 
nerves on each side prominent, curved-ascending, near the margin 
arched anastomosing, the lower pair usually forming an acute angle, 
ascending and more distally separated from the others, the transverse 
tertiary nerves thin but prominent, the minor veins forming a promi- 
nulous conspicuous reticulum, the costa and the principal lateral 
nerves covered when young, mainly towards the base, by a floccose 
deciduous indument of ochraceous, entangled, stellate hairs, in time 
becoming glabrous, shining, marked with minute callose, reddish, 
sparse dots, the tertiary nerves also glabrous, the minor veins, reticu- 
lum, and areoles covered by minute, white, stellate hairs forming an 
appressed, whitish tomentum. 
Inflorescences very small, axillary or in exfoliated, thin branchlets, 
the axis and branchlets of the cymes very reduced, ochraceous- 
ferruginous, lanuginose-tomentose; peduncles 2-7 mm. long, 3-bracteo- 
late at apex;'pedicels 3-8 mm. long, somewhat thicker; bracteoles 
560 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
linear, 1.5-2 mm, long; buds globose, also ochraceous-lanuginose and 
woolly-tomentose. 
Sepals thick, ovate, subacute, slightly involute-marginate, about 
6-7 mm. long, 3-4 mm. broad, united 1 mm. at base, rosy or reddish 
inside, subglabrous, glandular at base, 3-5 nerved, the margin mi- 
nutely whitish tomentose, rather woolly outside, stellate-tomentose, 
spreading in an thesis; petal-hoods yellowish or reddish obovate, 
rounded cucullate and slightly retuse at apex, with 7 prominent nerves 
and copious, spreading, minute hairs inside, subglabrous with few, 
slender, flexuous hairs outside, 3-3.5 mm. long, 2.5 mm. broad; petal- 
lamina deep red, rather thick, suborbicular, often retuse at apex, 
abruptly contracted into a pedicel at base, pilose at margin, and with 
very sparse, flexuous, slender hairs on the inner side, 3.5 mm. long, 
4 mm. broad; pedicel red, 0.6 mm. wide with sparse, slender hairs, 
2.5 mm. long. 
Androecium tube about 1.5 m. high, glabrous; staminodes laminar, 
thick-inembranaceous, deep red, oblong-elliptic, rounded and often 
emarginate at apex, attenuate near the base, with a conspicuous medial 
nerve and thin, flexuous, subspreading hairs distributed throughout, 
especially on the margins, 5-6 mm. long, 2.5-3 mm. broad; filaments 
thick, glabrous, 1.5 mm. long, shortly 3-furcate, 3-antherif ero us; anther 
lobes ellipsoid, 0,4-0.5 mm. long; ovary ovoid, 1.5 mm. long, densely 
tomentose-hirsute; styles glabrous, 5 mm. long, united only at base. 
Fruit obovoid-ellipsoid, rounded at apex, contracted at base, 
greenish, when ripe brown yellowish, 5-7 cm, long, 3-4 cm. broad, 
the pericarp thin-coriaceous densely covered with acute, hard warts 
and sparse stellate hairs, when dry about 1,5 mm. thick; seeds 16 
mm, long, 9 mm. broad; germination hypogeous. 
Common names.—Cabega de urubti is the most common and 
widespread. Also, the following have been locally recorded: 
copu-ai, cupu-curda, cupurana, cacao de macao, urubu-acain, 
cab eg a de Umbti. 
Win-che6k (Puinave), Infrida-Guaviare; ma-oo-hee-rGe (Ka- 
buyari) (Rio Cananari) (Angl. Col. Cocoa Exped., Baker 1952). 
Distribution.—Spread throughout the western part of the Amazon 
basin, on elevated ground and humid, fertile soil of rain forests. In 
Brazil it is frequent in the western half of Amazonia, the easternmost 
localities known being Teff6, on Rio Solimoes and Rio Jau, a tributary 
of the Rio Negro (Duche). The Anglo-Colombian Cacao Expedition 
found it (although not abundantly) in the rivers Cagu&n, upper and 
lower Caquetfi, and the Putumayo (Report, 1954, 12, 13). In Peru 
goes as far as the lower parts of Rio Huallaga and Rio Ucayali, Rio 
Itaya, and also Putumayo. 
CUATRE CASAS—CACAO AND ITS ALLIES 561 
m 
;i«W     •/.? 
c 
I * /■**■■ 
"I. U4,*,||i ' t  *  ■ t t*'«,-.1l1 »  ** ,,.+U, H, ■ 1 B-- v+- ■ fc7- !■■ ■■ i I ■ ""  . _ .  »* fc * H * I ■ * flk ■ ■ - p" 1 Ijfljk *J| ■■ -U J k"f 4+f ■■* | | P J ■ P+ w ■ k I L\ at, • * - M»4M 1 H11^ b !■» i *MP*+d ■ O . , ■• ■ ''jT" 1 1 , I I " \ % J +a* 'u.h.T''1 k» 
fe:-v"i\ i /- ■■-»/ .iv\ «;J-V ^    ^1" 
D 
Figure 40.—Detail of indumcnt on the underside of leaf in: a, Theobroma simiarum (Stand- 
ley 37377) from seedlings; b, T. simiarum (Pittier 7731), from adult branches; c, T. 
grandifiorum (Killip & Smith 30011); d, T. obovatum (Ducke 265). a X 20, b X 30, 
c and d X 40, 
562 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Tkeobroma obovatum is a rather small tree with a crowded, leafy 
crown, easy to recognize by its small, ellipsoid-obovoid fruits con- 
stricted at the neck, with thin, fragile, granulate or verrucose pericarp, 
usually green but becoming brownish yellow when ripe. Also 
characteristic are its light, papery, asymmetrical, oblong-obovate 
leaves with a close, white or cinereous monotrichous tomentum 
beneath; when very young the upper side, the principal nerves as 
well as the petioles, stipules, and twigs, are covered with an ochraceous 
brown or orange, easily removed down, composed of long, deciduous, 
stellate hairs. The fruit when ripe separates from the pedicel and 
falls. 
Schumann did not know the species, for it is lacking in his her- 
barium; he included the name T. obovatum as a synonym of T. sub- 
incanum in his treatment in the Flora Brasiliensis (p. 77), 
Huber knew the species well, although he called it T. sylvestre, 
confusing and misinterpreting Aublets figure of Cacao sylvestris. 
Ducke also knew T. obovatum and its distribution very well, but he 
thought that T. sylvestre Mart, was a synonym. 
COLOMBIA: Putumayo: Rfo Caucayd, Rio Leguizamo, Laguna Primavera; 
tree 15 m., 3 IV 1953, Holliday & Cope T/90 (COL, TRIN, US). Ibidem; tree 
10 m., flowering, on elevated land above river, 5 IV 1953, Holliday & Cope T/95 
(COL, TRIN, U, US). 
Caqubta: Rio Cagu&n, camp 4, 27 IV 1953, Holliday & Cope T/119 (COL, 
TRIN, US). Ibidem; found on sloping land, 30 ft., above river, tree 6 m,, 
jorquettes arising symmetrically, 26 IV 1953, Holliday & Cope T/114 (COL, 
TRIN, US). 
Amazonas: Rio Caquetd, La Pedrera, river level; tree 10 m., native in forest 
on the riverbank, 7 X 1952, Baker & Cope 30 {COL, P, TRIN, US). Ibidem; 
tree 7-8 m. in forest river-bank, presumed native, jorquettes regularly of three 
branches growth continuing from above, young twigs and leaves with caducous 
fuzz, flowers pale crimson, 5 X 1952, Baker tfe Cope 27 (COL, F, TRIN, US). 
Ibidem; large tree 30 ft. high; calyx lobes light outside and pinkish inside, petals 
dark purplish red, diameter 9-10 inches, 5 X 1952, SckuUes & Cabrera 17775 (US). 
Rio Caquetd, Remolino; tree 6-8 m., no flowers but had two ripe pods, 2 V 1953, 
Holliday & Cope T/123 (COL, TRIN, US). Trapecio Amaz6nico, Loretoyacu 
River, 100 m. alt., XI 1945, SckuUes 6921 (F). 
BRAZIL: ParA: Bel6m, Jardim Bot&nieo do Museu Goeldi; small tree, "cacau 
c&beQa de urubti," cultivated, 11 VIII 1942, Archer 7537 (USDA, IAN). Ibidem; 
arvore no. 482, 22 XII 1958, Cavalcante 339 (MG, US). Ibidem, Pires & Black, 
s.n. (BH), 743 (IAN). Ibidem (Rio Puriis, loco dicto Bom Logar oriuntur, J. 
Huber anno 1904 accedit); arbor parva floribus atrorubris, fructus maturitate 
flavidia, "cabc$a de urubu," 4 II 1926, Ducke 21044 (G, K, P, U, US). Ibidem, 
Ducke 265 (A, F, K, MO, S, US). Bel6in, cultivado in Instituto Agron6mico 
do Norte, "cabega dc urubii," Pires, Nilo, <fc Silva 4339 (IAN, UC, US). 
Amazonas: San Antonio do Iga, matta, 27 IX 1906, Dueke 7704 (MG). 
On the Rio Negro, Sckomburgk 870 p.p. (L), Municipality Sfio Paulo de Olivenca, 
m?ar Pal mares; tree 35 ft. high, trunk 4 inches diam., terra firma highland, 11 
IX-26 X 1936, Krukoff 8275 (A, BM, FE, F, G, K, MICH, MO, P, S, U, US, 
USDA, Y). Basin Rio Madeira, Municipality Humayta, near Tres Caaas, on 
CUATRECASAS—CACAO AND ITS ALLIES 563 
varzea land, shrub 25 ft. high, "cabe^a de umbti," 14 IX-11 X 1934, Krukoff 
6263 (A, BM, F, GOET, K, LB, MICH, MO, S, U, US, Y). "Amazonas Ega," 
Poeppig 2746 p.p. (LE). "Brasilia in sylvis circum Ega, 1831," Poeppig, s.n. (WU). 
Amazonas; Boeca et TeffG, matta, "cabega de urubd," 27 X 1904, Ducke 6823 
(BM, G, MG, P, US). Teff6, matta virgen; tree, flower white, "copu-ai," 22IX 
1947, Black 47-1496 (IAN). Igarap6 Jandiatuba, lowland, border of the 
river, 14 I 1949, FnSea 23926 (US, IAN). Arredodes de Fonte B6a, terra firma 
alta; arvore media, "cupurana," "cupu-curua," 12 IV 1945, Fr6es, 20646 (F, K, 
IAN, USD A). Rio Jurud, Jurud Miry; Baum 3-9 m., Blumen dunkclpurpurn, 
VIII 1901, Ule 5637 (G, HBG, K, MG, L). Purtis, Monte Verde, "cacao de 
macaco," II 1904, Goeldi 4226 (MG). Rio Acre: Antimari, matta, 31 III 1904, 
Huber 4295 (BM, G, MG, P, US). Upper Amazon, Paranary; low tree, petals 
purple, stamens yellowish, staminodia yellow, "urubd-acaim," 20 X 1924, Traill 
61 (K, P). 
Acre: Basin Rio Purtis, near mouth of Rio Macauh&n (tributary of Rio 
Yaco), Lat. 9°20' S, Long. 69° W, on terra firma; tree 35 ft. high, 9 VIII 1933, 
Krukoff 5388 (A, G, K, MICH, MO, S, U, US). Ibidem; shrub 40 feet high, 
3 IX 1933, Krukoff 5759 (A, BM, F, G, K, MICH, MO, S, U, US). 
Mato Grosso: Machado River region, source of the Jatuarana River; tree 
45 ft. high in terra firma, "cupuarana," XII 1931, Krukoff 1668 (A, BM, G, K, 
F, MICH, MO, S, U, UC). 
Guaporb: Porto Velho, km. 8, matta on elevated ground; small tree, 17 
VI 1952, Silva 155 (IAN). 
PERU: Loreto: Along Rio Itaya, Rfo Masana, 8 V 1929, LI. Williams 161 
(F). Forest of Rfo Itaya, 3 V 1929, LI. Williams 230 (F, US). Lower Rfo 
Huallaga, Puerto Arturo, Yurimaguas, 155-210 m., medium-sized tree, forest, 
20 XI 1929, LI. Williams 5268 (F, US). Balaa-puerto, 220 m., forest; tree 4 m., 
flowers wine red, IV 1933 Klug 2983 (A, BM, G, GH, K, F, MO, S, US). May- 
nas, in sylvis circum Yurimaguas, 1831, Poeppig 1845 (G). Maynas 1831, 
Poeppig 2352 p.p. (lectotype, WU; isosyntypes, F, GOET, G, LE, P), Stromgebiet 
des Ucayali, von 10° S bis zur Mundung, 1923, Tessmann 3433 (G, S). Rain 
forest of the Amazon basin, 230 m,, 40 km. south of Pucallpa, 24 VII 1957, 
Ellenberg 2551 (L). Bank of Rio Putumayo, opposite Puerto Leguizamo; sucker 
of large flowering tree, leaves rather large, 7 IV 53, Holliday <fc Cope T/98 (COL, 
TRIN, US). 
TRINIDAD (cult.): Imperial College of Tropical Agriculture, River State 
Diego Martfnez, Field 19; primary branching ternate; growing pseudoapical; 
branchlets and twigs ochraceous lanate-tomentose, leaves thin, flexible, very pale, 
with costa and secondary nerves yellow green, with woolly, floccose, deciduous 
tomentum beneath, green above, 31 VIII 1961, Cuatrecasaa, Cope, & Bartley 
25783T (US). Ibidem; trunk 15 cm., bark lenticellate-granulate; branching 
ternate; terminal branchlets light brown or brownish ochraceous, lanate-tomen- 
tose, crown very branched and leafy, the lower branches horizontal, the upper 
ones ascending, young, tender, terminal leaves hanging, light yellowish green, 
1 IX 1961, Cuatrecasas & Cope 25788 (US). 
17. Theobroma subincanum Mart. 
Figures 29, 33, 35, 39, 41; Map 11; Plates 9, 10, 11 
Theobroma subincanum Mart, in Buchner, Repert. Pharm. 35:23. 1830; 
Linnaea Litt. Bericht, 32. 1831; Bernoulli (1869) 13; Schumann in Mart. 
(1886) 77; Jumelle (1899) 27 (in part); Huber (1906a) 274; Ducke (1925) 
132; (1940) 272, pL 4> fig- U Addison & Tavares (1951) 25, pi. 8, fig. 1, 
564 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Map 11,—Geographical distribution of Thtobroma subincanum # and of its vicariants at 
the western side of the Andes, T. hylaeum A and T. nemoralt Q. 
pl• 4>fig- A, pl. 11, fig. 3; Ducko (1953) 10; Cuatrecasas (1956) 699; Baker, 
Cope, & al. (1954) 12, fig. IS. 
Cacao sylvestris Aubl. PI. Guian. 2:687, pl. 276. 1775. 
Cacao guianensis Aubl. Pl. Guian. 2:684. 1775, pro parte (tantum folia). 
Tkeobroma sylvestris (Aubl.) Don, Hist. Diehl. Pl. 1:622. 1831, non Mart. 
1830; Chevalier (1946) 279; Lerade (1952) 380; Le6n, (1960) 322, 321, fig. 
Theobroma ferruginea Bernoulli, Uebers. Art. Theobroma 13. 1869. 
Theobroma alba Ruiz & Pav6n, Fl. Peruv. Chil. 6, pl. 68t ined. 
Types.—Amazonas, Brazil, Martins. French Guiana, Aublet (of 
Cacao sylvestris). Peru, Ruiz db Pavon (of T. ferruginea). 
Medium-sized tree commonly 6-12 m. tall, at times up to 20 m. high, 
the trunk 15-20 (-30) cm. in diameter, with gray, almost smooth 
bark, older bark rugose-rimose,' reddish within, the wood whitish, 
darker toward the center; growth pseudo apical; primary branches 
ternate, grayish, spreading; juvenile branchlets covered by a dense 
ferruginous tomentum of stellate hairs, when older glabrescent, pale 
brownish or brown, somewhat rugose, rimose-reticulate; stipules 
narrowly linear, densely ferruginous-tomentose, 5-7 mm. long, 1 mm. 
wide, soon deciduous. 
Leaves firmly coriaceous, rather thick and large; petiole robust, 
subterete, densely and appressed ferruginous-tomentose, 8-15 mm. 
long; lamina elliptic-oblong or subobovate-elliptic-oblong, very little 
CUATRECASAS—CACAO AND ITS ALLIES 565 
attenuate to the base, slightly unequal, rounded or very obtuse, 
emarginate or rarely cordate at base, somewhat narrowed or rounded 
and abruptly acuminate at apex, sometimes blunt, entire, or near the 
apex dentate-sinuate, 16-40 cm. long, 5-20 cm. broad, the acumen 
acute, 1-3 cm. long, when very young ferruginous-tomentose through- 
out, but soon glabrescent above, when adult glabrous above, green, 
somewhat brownish olivaceous when dry, the costa and the lateral 
nerves depressed, filiform, the lesser veins obsolete, cinereous beneath, 
the veins more or less tawny or ferruginous, the costa thick, very 
prominent, the 9 or 10 pairs of secondary nerves very prominent, 
subascending, thinner near the margin, decurrent, the superior arched, 
anastomosing, the basal pair often straighter and forming a more acute 
angle, the transverse tertiary nerves prominent, the minor ones and 
small veins prominulous, minutely reticulate, the midrib, major nerves, 
and reticulum more or less densely covered by mediocre, reddish or 
tawny stellate hairs, the areoles between the veins with a dense 
whitish indument of very small, delicate, intricate, stellate hairs. 
Inflorescence small, few-flowered, axillary or extra-axillary on leafy 
branches; cymes with 3-9 fasciculate branchlets, usually 1-3-flowered; 
peduncles 2-8 mm. long, with 3 bracteoles at apex, the bracteoles 
subulate, about 3 mm. long, deciduous; pedicels 3-6 mm. long, 
thicker than the peduncle; buds ovoid-globose; sepals thick, carnose, 
ovate, acute or subacute, densely stellate-tomentose outside, ferru- 
ginous, the margin minutely whitish tomentose, shining inside, 
purplish or red, subglabrous with minute, crowded, oblong-capitate, 
glandular hairs at base, near the margin slightly pubescent, 8-9 mm. 
long, 3-4 mm. broad, united at base for 2 mm., subpatulous. 
Petal-hoods thick-membranaceous, pale yellow and red striate, 
obovate, rounded-cucullate at apex, slightly emarginate, 7-nerved, 
inside minutely hirtulous, glabrous outside except for the puberulous 
margin, 3-3.5 mm. long, 2-2.4 mm. broad; petal-lamina pedicellate, 
carnose, thick, rigid, red, suborbicular, 2-2.5 mm. long, 2.2—4 mm. 
broad, with slightly retuse apex, slightly pilose at margin, the hairs 
very slender, flexuous; pedicel 2 mm. long, compressed, pilose. 
Androecium tube 1.5-1.7 mm. long, glabrous; staminodes laminar, 
red, lanceolate-oblong, acute or subacute, with marked midrib, sub- 
glabrous with sparse flexuous hairs at margin, 6-7.5 mm. long, 2 mm. 
broad; filaments rather thick, glabrous, about 1.5 mm. long, arched, 
very shortly 3-furcate, 3-antheriferous, the anther cells ellipsoid, 
about 0.5 mm. long; ovary ovoid-oblong, 1.3 mm. long, glabrous, with 
very sparse, minute, granulate dots; styles 1.5 mm. long, connivent. 
Fruit ellipsoid, light green, tawny or orange at maturity, smooth, 
oblong-ellipsoid or obovate-ellipsoid, rounded at apex, often more 
or less narrowed at base, 7.5-11.5 cm. long, 5-6.6 cm. broad; pericarp 
566 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
coriaceous, rigid, hard, 3-4 mm. thick, the woody epicarp 1-2 mm. 
thick, covered by a brown, thin, appressed, stellate-tomentose in- 
dument; seeds ellipsoid-oblong or ovate-ellipsoid, 1.8-2,3 cm. long, 
12-16 mm. broad, 8-11 mm. thick, the surrounding pulp rather 
slightly sweet, scentless, white, becoming yellowish; cotyledons white; 
fruiting peduncle 1-1.5 cm. long, 0.5-1.0 cm. thick; germination 
hypogeous. 
Common names.—The commonest names in Brazilian Amazonia 
are cupul and cupual. Other names or other ways of spelling and 
pronouncing the former are: copui, copuaf, cupuhy, cupuahy, 
cupuy do Igap<5, cupuarana, cupti do matta, cupfi-assuy, cupfi-assti- 
rana. In Colombia, Venezuela, and Peru this species is usually 
called cacao do monte or cacao silvestre, and also cacao rana (Orinoco 
valley), yurac-cacao (Yurimaguas), uchpa-caeao, cacao-ceniza (Peru), 
cacao bianco (Peru, Ruiz db Pavon). Indian names recorded are: 
abekard (Makunat Vaup4s, Garcia Barriga), padama (Arekuna, 
Venezuela, cumala (Peru). The Anglo-Colombian Cocoa Expedition 
(Baker, 1952) recorded the following indigenous names: Win-che&k 
(Puinave), Inirida-Guaviarc; bawk (Maku), Piraparana-Taraira; 
poo-hoo (Barasana), Upper Piraparana; a-ba-ka-ra {Makuna), Lower 
Piraparani Popeyaca; mali-we-re (Yukuna)f Miritiparand; no-t6r- 
ree-ka (Tanimuka), Guacaya; too-soo (Yauna), Lower Piraparand,; 
ma-oo-hee-re6 (Kabuyari), Canarari; wa-k6 (Kubeo), Cuduyari; 
wah-pek-Ia (Tukano), Papuri; a-so-ya-ee (Piratapuya), PapuH; 
wa-be-ga-ra (Desano), Papuri; wa-be-ka-ra (Siriano), Paca; ma-w6- 
roo-da (Kuripaka), GuainJa. 
Uses.—Although this species gives an acceptable chocolate it 
is practically never used by the natives. The slightly sweet and 
scentless pulp is occasionally eaten or sucked; it is very much sought 
by animals, especially monkeys. 
Distribution.—Widespread throughout Amazonia from Parfi to 
the most western tributaries of the Amazon River, the upper Orinoco 
range, and the Venezuelan and French Guayanas; frequent in the 
shade of the Hylaean rain forests, in noninundatable lowlands, often 
on rich and humiferous soil but ascending small hills on sandy grounds, 
along creeks and small rivers. Theobroma subincanum is the species 
of most frequent and abundant occurrence and with the broadest 
area of distribution, other than T. cacao. 
COLOMBIA: Meta: Acacias, Canaima, farm 350 m. alt., cultivated, 18 XI 
1951, Patino 22 (F). Sierra de la Macarena, trail from Rfo Gufijar to Cafio 
Guapayita, Cafio Yerli, 500-600 m. alt.; tree about 35 ft. tall, flowers deep red, 
fruit ripening brown, leaves rusty beneath, 20-28 XII 1950, Idrobo Schulies 
776 (COL, IAN). 
Putumayo: Rfo Caucayd, Laguna Primavera on Rfo Legufzamo; tree 18 m., 
symmetrical jorquette, 3 IV 1953, Holliday & Cope T/91 (COL, TRIN", US). 
CUATRECASAS—CACAO AND ITS ALLIES 567 
A
 B 
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31 
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r <v:-;V v|; 
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"*k 1 ' I J ■ Cla AlaJailltl II II III i t 7 I Jf" ■ " hi IK" 'W f #«*,"*#+* < ill | fl+l « 1 I i; : * 'mi ii 
D 
Figure 41.—Detail of indument on the underside of leaf in: a, Theobroma subincanum 
(Cuatr. 7277); b, T. sinuosum (Pavon, s.n.)i c, 7*. nemoralt (Cuatr. 21921); d, T. mam- 
mosum (Leon 1363). A, c, and D X 25, b X 10. 
Vicinity of Mocoa; tree in forest, 15 m. high, no fruits or flowers, 17 III 1953, 
Holliday & Cope T/81 (COL, TRIN, US). 
680-695—64 13 
568 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
CaquetA: Rfo Cagu&n, camp 4; tree 10 m.t 26 IV 1953, HoUiday & Cope 
T/116 (COL, TRIN, US). Upper Rfo Cagu&n; small trees, 10IV 1953, HoUiday 
& Cope T/101 (COL, TRIN, US), 13 IV 1953, HoUiday & Cope T/103 (COL, 
TRIN, US). Solana, 8 km. SE. of Tres Esquinas, on Rio Caquetd, below mouth 
of Rfo Ortegiiaza, 200 m. alt., wet tropical forest; tree 15 m. high, 15 cm. thick, 
bark gray, smooth, with lichen patches and bryophyte patches, long, ellipsoidal 
orange-brown fruits 10 x 6-6.5 cm., edible, "cacao silvestre," 4 VI 1945, Little 
& Little 9544 (F, US) (fruit US No. 1096). Municipality of Florencia, site San 
Luis, right margin of Pescado River, "cacao silvestre" 5 VI 1942, Rangkel 195 
(in part) (COL). 
VAUpfis: Cafio Mayabo, near San Felipe, river level; only flowers and old 
fruits, no young or ripe fruit at this season, 27 X1952, Baker 33 (COL, F, TRIN, 
US). Mitfi: Cafio ParanA-Midf, 200 m. alt.; tree 5 m., 19 X 1939, Cuatrecasas 
7277 (F, US, COL). Cerro de Mitu, 380 m. alt.; small tree with bending, fruiting 
branches, fruits 9.3 X 6 cm., 17 IX 1939, Cuatrecasas 6890 (COL, F, US). Mitfi 
and vicinity, 280 m. alt.; tree, cultivated, 22-30 VI 1958, Garcia Barriga, Schultes 
& Blohm 16064 (COL). Ibidem; tree 5 m. with spreading branches, flowers red, 
10-13 XI 1952, Garcia Barriga 15139 (COL). Cerro de Circasia, 300 m. alt., 
base of hill 10 X 1939, fruiting tree, 10 X 1939, Cuatrecasas 7178A (COL, F, US). 
Rio Barbas, river level, native in the forest, 28 X1952, HoUiday 43 (COL, F, TRIN, 
US). Rfo Cuduyarf, tributary of Rio Vaupfe, Yacara, middle and lower course, 
highland, ±700-800 feet, 9 XI 1952, Schultes, Baker, & Cabrera 18552 (US). Rfo 
Cuduyarf, river level, showing jorquette of 3 branches, 16 X 1952, Baker & Cope 31 
(F). Ibidem; young red flowers, Baker, Bartley, & HoUiday 31 (COL, TRIN). Rfo 
Cuduyarf, Pakwativa Mission, river level, heavily infested with Marasmius 
perniciosits, 19 X 1952, Baker & Cope 32 (COL, F, TRIN, US). Ibidem; flowers 
distinctly paler in color, Baker, Bartley, & HoUiday 32a (TRIN). Rfo Apaporis, 
Soratama, above mouth of Rfo Cananarf, Cafio Surruco, 900 feet alt., highland; 
small tree, leaves rusty beneath, fruit rusty brown, ovoid, 30 I 1952, Schultes <fc 
Cabrera 15116 (US). Rfo Apaporis, Rio Jinogojfi, river level; tree 40-50 ft., 
native in forest or river bank, 13 IX 1952, Baker <fc Cope 12 (TRIN). Jinogojd, 
growing just above flood level with numerous ellipsoid pods, Baker & Cope 7 
(COL, TRIN). Ibidem; river level, tree 16 m., 20 cm. diam. at base, a long, tall 
thin pole, without obvious jorquettes, native in forest, 23 VIII1952, Baker <fc Cope 
3 (COL, F, TRIN, US). Ibidem, 250 m. alt.; tree 15 m., large, coriaceous leaves, 
brownish fruits 7.5 X 4.5 cm., 25, 26 VIII 1952, Garcia-Barriga 14224 (COL, US). 
Rfo Piraparanfi, cultivated tree, 21 VIII1952, Schtdtes & I. Cabrera 17005 (AMES). 
Rfo PiraparanA, 250 m. alt.; tree 3 m., red petals, sepals red above, 22-26 VIII 
1952, Garcia-Barriga 14203 (COL, US), 14253 (COL). Rfo PiraparanA, near 
confluence with Rfo Apaporis, river level; small tree 3 m., in Indian garden, 
24 VIII1952, Baker & Cope 4 (COL, F, TRIN, US). Rfo Infrida, near Morichal, 
near the mouth of Rfo Papunand, 200 m. alt.; tree 10 m., 14 II 1953, Fern&ndez 
2275 (COL, US). Rfo Infrida, Raudal, 300 m. alt., 3 II1953, Bartley & HoUiday 
T/71 (COL, TRIN, U, US). Rfo Infrida, Santa Rosa; tree 12 m., 25 I 1953, 
Bartley & HoUiday T/68 (COL, TRIN, U, US). Rfo Infrida, right bank below 
Cafio Caribe (5 hours above Morichal); tree about 10 m., no flowers, one small 
fruit, 2211953, Bartley & HoUiday T/65 (COL, E, TRIN). Rfo Infrida, Morichal, 
30 m. alt., in forest, small fruits, no flowers now, 8 II 1953, Bartley & HoUiday 
T/72 (COL, TRIN, US). Rfo Infrida, affluent Papunandi, 300 m., red leaf flush, 
nearly mature pod 11.5 X 6 cm., fruit pedicel 2 cm. long, fruit green with brown 
pubescence, pericarp woody, pulp white, cotyledons white, convolute, 18 II 1953, 
Bartley & HoUiday T/75 (COL, TRIN, US). 
CUATRECASAS—CACAO AND ITS ALLIES 569 
Amazon as: Rfo Caqueti, La Pedrera, river level; tree 45-50 ft., native in 
forest, 29 IX 1952, Baker & Cope 25 (COL, F, TRIN, US). La Pedrera, river 
level, native tree in forest, 26 IX 1952, Baker & Cope 21 (TRIN), Rfo Caqueta, 
Remolino, leaves from young tree, 2 V 1953, Hottiday & Cope T/124 (COL, 
TRIN, US). Rfo Apaporis, near mouth of Rfo Cananarf; tree 45 feet tall in 
forest, fruit rust colored, "cacao de monte," III 1951, Schultes 12104 (COL, US). 
Rfo Apaporis, between Rfo Pacoa and Rio Cananarf, Soratama, 250 m. alt.; weak 
tree, leaves rusty beneath, flowers red, 26 IX 1951, Sehidtes <fe Cabrera 14140 
(US). Rfo Miritiparand, near varadero to Rfo Apaporis, river level; tree 50 ft., 
native in forest, 15 IX 1952, Baker & Cope 13 (COL, TRIN). Trapecio Ama- 
z<5nico, Loretoyacu River, 100 m. alt., IX 1946, Schultes 8385 (AMES, F). Leticia, 
100 m. alt.; bark rough, cracked, light gray, flowers red, 20 IX 1945, Schultes 
6536 (F). Leticia, forest; flowers pale red, tree 5 m., 22 VIII 1946, Black <fe 
Schultes 46-61 (USDA). 
VENEZUELA: Bolivae: Mount Duida, 500 m. alt.; small tree, VIII 1928-1V 
1929, Tale 944 (NY, US). Caronf, rain forests of the Icaburu valley, 440 m. 
alt.; tree 15 m., fruit large and tasty, "padami" (Arekuna), XI 1947, Cardona 
2379 (US, VEN). 
Amazon as: Alto Rfo Orinoco, TamanA, 121 m. alt.; medium-sized tree (10-12 
m.) with few branches, flowers purplish or red on the branchlets, trunk bent, up 
to 40 cm. diam. with no branches in 2-5 m., bark gray, inside light brown, wood 
pale brownish; "cacao-rana," LL Williams 15204 (A, F, G, US, VEN). Rfo 
Guainfa, Maroa, river level, with one ripe pod (wall mealy, smooth, thin and 
brittle), seeds extracted and sent to Trinidad, Baker 38 (COL, F, TRIN, US). 
FRENCH GUIANA: Aublet s.n. (part of syntype of Cacao guianensis Aublet, 
BM, Photo, Mo. Bot. Card. no. 4028). "Guyane Francaise," Poiieau, s.n. 1819- 
1821 (G). French Guiana, "Cacao sylvestris Aublet," Aublet (syntype, BM, Photo 
Mo. Bot. Gard. no. 4029). 
BRAZIL: Amazonas: Rio Negro, Barcellos, "cupuhy," 25 VI 1905, Ducke 7200 
(MG). Rio Negro, Rio Caure, Igarap6 Miritf; tree 5 m., 15 cm. diam., red 
flowers, in rather lowland high forest, VII 1948, Fries 23343 (IAN, P). Rio 
Negro, Porto Cabary, "cupuhy," 4 XII 1945, Frdes 21482 (IAN, NY, US DA). 
"Prov, Rio Negro Martius Iter. Brasilienses 325," Martins [898] (M) (Photo 
F. M. 19645). "Prov. Rio Negro, Martius Iter. Bras. 325," Martius [872,873 p. p., 
894, 895, 896, 897, 899, 900] (M). "Rio Negro Dr. Martius Iter Brasil. 325, 
Theobroma subincanum," Martius [893] (M) (Photo F. M. 40704). Rio Vaup&: 
Jauratl, inundatable forest, 17 X 1945, Frdes 21162 (IAN, NY, USD A). San 
Antonio de Iga, forest; tree 10 m. "differt a T. ferrugineo Bern, foliis majoribus 
latioribus floribus majoribus," Ducke 7679 (MG, F) (Photo F. M. 7679). Mu- 
nicipality of Sad Paulo de Olivenga, near Palmares; tree 40 ft. high, trunk 5 
inches diam., high land, "cupuarana," 11 IX-26 X 1936, Krukoff 8226 (A, BM, 
F, G, GB, K, LE, MO, NY, P, U, US). Basin of Rio Madeira, Municipality 
Humayta, near Livramento, on Rio Livramento, terra firma; tree 60 ft. high, 
"cupuarana," 12 X-6 XI 1934, Krukoff 7016 (A, F, G, K, MICH, MO, S, U, 
US, WU). Rio Madeira, Varadouro do Morcego, 31 VIII 23, Kuhlman 18110 
(U). Manaos, mata; arvore, tree 30 m. alt., 15 cm. diam., "cupuhy," 17II1945, 
Frdes 20518 (IAN, K, NY, USDA). Manaos, Mata do Aleixo, "cupuhy," 16 
III 1945, Frdes 20555 (F, IAN, USDA). Manaos Aurora Fazenda, 15 m., 
Urwald, "cupd do Matto," 28 VIII 1921, von Luetzelburg 22079 (M). Without 
locality Spruce 97 (K). 
AuapX: Rio Amaparf, Serra do Navio, slopes of Curuca Ore Body, down 
to Igarape Sentinela; occasional tree 6 m. tall; fruit brown, 9 XI 1954, Cowan 
38186 (NY, US). Lower slopes of Observatorio Ore Body, heavily forested hills, 
570 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
70-300 m. alt.; tree 10 m. tall, fruit brown, 8 XI1954, Cowan 38164 (NY). Missao 
do Servico Florestal no T. Amap&, IX 1955, Miranda Bastos 68 (IAN). 
PabA: Bel&n, "cupuahy," VI1896, Huber 162 (BM, G, MG, P, (US). Bel&n, 
Bosques Rodriguez Alves, 1 VIII 1044, A. Silva 317 (IAN, USDA). Bel6m, 
Horto do Museo Goeldi; tree 481, small tree, flowers brown yellow, 22 XII 1958, 
Cavalcanle 938 (MG, US)- Bellm, Bosque municipal; tree 30 feet high, sparsely 
branched, 4 VIII 1942, Archer 7517 (IAN, USDA). Beldm, south forest of the 
Instituto Agrontfmico do Norte; large tree, contents of fruit eaten by birds, 
"cupu-assuy," 16 XI 1942, Archer 7820 (IAN, USDA). Bel£m, along roads on 
lands of Inst. Agr. do Norte; large tree, flowers rose color, "cupu-assu-rana," A. 
Silva 237 (IAN). E. F. Bragan$a, Joao Coelho; tree 8 m., 14 III 1947, Pires 
<fc Black 1414 (IAN). Region of Igarapfi Pitor6; tree 10 m., flowers red, 19 IX 
1958, Fries 34663 (IAN). Taperinha, near Santarlm, bushed river margins of 
Igarap6 Assti, "cupuy do Igap6," 23 VIII 1927, Ginzberger & Hagmann 801 
(F, WU). Rio Tapaj6z, Cachoeira do Mangabal, "beira de assahyzal," "cupuy," 
6 IX 1916, Dueke 16464 (BM, MG). Matta do Alto Ariramba, ' cupuy" 7 X 
1913, Ducke 14925 (MG). Rio Puriis, Monte Verde, "cupuahy," II 1904, 
Goeldi 4225 (MG). Paranary, upper Amazon, "cupua-i," 20 X 1874, Traill 59 
(K, P). 
Guapob6: Porto Velho, km. 8, in high forest on firm land; tree about 30 
m., "cupul," 17 VI 1952, J. F. Silva 143 (IAN). 
Rio de Janeiro: Quinta de S. Christovao; small tree planted by Riedel 
probably originally from Amazon basin, "cupual," 16 II 1876, Glaziou 9633a (P). 
PERU: Loreto: Mishuyaco, near Iquitos, 100 m. alt., forest; tree 6 m. 
high, flowers dark red, "wild cacao," X IX 1929, Klug 87 (F, NY, US). Ibidem, 
II—III 1930; tree 8 m. high, flowers wine red on branches, Klug 857 (F, US). 
Alto Itaya, 145 m., in forest of Paraiso, "cumald," 30 IX 1929, LI. Williams 
3254 (A, F, G, S, US). Upper Rfo Nanay, Santa Ana, "uchpa-cacao," 7 VII 
1929, LI. Williams 1233 (F, S, US). Rfo Nanay, Tierra Doble, deep forest, 
"Campamento balatero Lira Dabu," 8 VI 1929, LI. Williams 1076 (US, F, S). 
"Peru," Ruiz & Pavdn, "Theobroma alba R. & P." (ined.), "Theobroma ferruginea 
Bern." (K, BM). Ibidem, Herb. Pavdn 201 (G). "Theobroma alba" Rivero 
1836 (P). Tessmann, s.n., NY-3717; probably the same as the Tessmann 4115, 
cited by Mildbraed (Notizbl. 11:139. 1931) as T.ferrugineum and collected near 
Pongo de Manseriche, 160 m. alt., tree 19 m. tall, 21 cm. diam., "pako-kakao" 
(Dunkcl Kakao), 23 IX 1924. 
HuAnuco: Pro v. Hu&nuco, Tingo-Marla, forest, tree about 25 m. high, 
sepals olivaceous yellow without, more or less red within, petals yellowish gray, 
31 VIII 1940 Asplund 13410 (S). Tingo Maria, forest; tree 15 m., flowers dark 
red, 8 VIII 1940, Asplund 12911 (S). Maynas, Yurimaguas, "yurac-cacao" 
i.e., "Cacao album Peruvianum," March, 1831, Poeppig 2352 p.p. (GH, P, WU). 
TRINIDAD (cultivated): On grounds of the I.C.T.A., from seeds from 
Mitd Vaup^s region, Colombia; one branch damaged by Marasmiut pemiciosus. 
Cuatrecasas, Cope, & Bartley 25785 T (U.S.) 
18. Theobroma hylaeum Cuatr., sp. nov. Figures 25, 39; Map 11 
Arbor circa 10 m. alta, apicaliter crescens, ramis primariis ternatis, 
ramulis brunneia juvenilibus minute ferrugineo-tomentosis denique 
glabratis, cortice rugoso-fissurato; stipulae lineari-subulatae acutae 
vel subacutae ferrugineo-tomentosac, circa 6 ram. longae, 1 mm. 
latae. 
CUATRECASAS—CACAO AND ITS ALLIES 571 
Folia mediocriter coriacea, petiolo crassiusculo subtereti minute 
ferrugineo-tomentoso, 6-10 mm. longo; lamina elliptico-oblonga vel 
obovato-oblonga, basi symmetrica attenuata obtusa vel subobtusa, 
apice leviter attenuata interdum rotundata subite acuteque acumi- 
nata, margine Integra vel apicem versus leviter undulata, 12-20 cm. 
longa, 4-9 cm. lata, apiculo 6-15 mm. longo, supra in sicco viridi- 
brunnescens glabra, costa et nervis principalibus filiformi-impressis 
reliquis vix conspicuis, subtus cinereo-ochracea vel nervatione fer- 
ruginea, areolis venulisque cinereo-tomentosis pilis stellatis minutis 
albidis intricatis dense tectis et pilis stellatis crassioribus ferrugineis 
sparsis vel in nervis copiosis, costa crassa elevata, nervis secundariis 
7 vel 8 utroque latere eminentibus arcuato-ascendentibus, prope 
marginem tenuioribus anastomosantibus, nervis tertiaris transversis 
prominentibus r nervulis venulisque tenuioribus sed prominulis 
reticulatis. 
Inflorescentiae breves axillares cymis 1-3 floribus instructae, 
ramulis brevissimis tomentosis, pedunculis tenuibus erectis vel sub- 
flexuosis ad 12 mm. longis ferrugineo-tomentosis apice 3-bracetolatis 
et cum pedicello articulatis; bracteolae anguste lineares subacutae, 
3-6 mm. longae, 1-2 mm. latae, extus tomentosae; pedicelli erecti 
tomentosi pedunculis leviter crassiores, 5-6 mm. longi. 
Sepala crassiuscula ovato-oblonga acuta, basi ad 2 mm. coalita, 
patulo-reflexa, extus ochraceo-tomentosa, intus margine minutissime 
albido-tomentella excepta glabra, aurantiaca, basi ad insertionem 
pilis crassiusculis oblongis glandulosis annulum formantibus, 8-9 
mm longa, 4—5 mm. lata; petala cucullo aurantiaco obovoideo, basi 
angustato, apice rotundato, saccato-cucullato, 3 mm. longo, 2 mm. 
la to, extus glabro ruguloso, intus minute hispidulo, 7-nervato, pediculo 
2.5 mm. longo, parce puberulo, lamina obovato-s ubr homboidea 
crassa brunneo-rubra, faciebus sparsis minutissimis pilis margine 
pilis flexuosis tenuibus praedita, 4-5 mm. longa, circa 4 mm. lata; 
androecium tubo circa 1.5 mm. longo; staminodia petaloidea aestiva- 
tione reflcxa in anthesin curvato-patula, brunneo-rubra obovato- 
oblonga apice rotundata vel leviter retusa, 5-5.5 mm, longa, 2.2 mm. 
lata, minute pilosula; filament a fertilia 1 mm. longa; antherae 3. 
brevissime pedicellatae; ovarium obovatum 1.2 mm. longum, 1 mm, 
la turn, crasse tomentosum; styli circa 1 mm. longi coaliti. 
Fructus coriaceus circa 7x4 cm., viridi-brunnescens, adpresse 
tomentosus laevis ellipsoideus, utrinquc rotundatus, epicarpio rigido 
lignoso in sicco 1 mm. crasso; pedunculus fructifer 1.5-1.8 cm. longus, 
0.8 cm. crassus. 
Type in the U.S. National Herbarium, No. 2028683, collected in 
the heavy rain forest around Villa Arteaga, northern region of An- 
572 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
tioquia, Colombia, at 200 meters altitude, August 14, 1948, by J. 
Araque and F. A. Barkley no. 18C745. 
Common names.—Chocolate de monte, cacao silvcstre. 
Distribution.—Presently known only from the Chocd region of 
northern Antioquia, Colombia, and probably also in Panama. 
COLOMBIA: Antioquia: Villa Arteaga, heavy wet forests at 200 m, alt.; 
tree 10 m., flowers orange, corona cream orange, stigma black brown, 14 VIII 
1948, Araque & Barkley 18C745 (holotype, US 2,028,683; isotype, US 2,028,684, 
COL). Ibidem; seedlings showing symmetrical branching in 3's, growth continu- 
ing from above the jorquette, 24 VII1953, Bariley & Holliday T 166 (K, TRIN, 
US). 
PANAMA: Col<5n: Along Rfo Fat<5, in forests or thickets 10-100 m. alt., 
VII 1911, Pittier 4194 (BM, GH, US) (probably; specimen sterile; identification 
needs confirmation by fertile specimens). 
Theobroma hylaeum is closely related to T. nemorale from which it 
differs essentially by the narrow-linear bracts and stipules, by the 
subrhomboid lamina, and by the narrower obovate-oblong, rounded 
and notched staminodia; furthermore, the peduncles and the pedicels 
are short, the pedicels shorter than the peduncles, and the fruit is 
harder, smaller and not constricted above the base. From T. subin- 
canum it differs by the shape of the petal-laminae and staminodes, 
by the tomentose ovary, by the smaller fruit and leaves, and by the 
venation, in which the minor veins are less conspicuous. 
19. Theobroma nemorale Cuatr. Figures 3, 41, 42; Map 11 
Theobroma nemorale Cuatr., Rev. Acad. Colomb. Cienc. 8:487, fig. 4. 1952; 
Baker, Cope, & al. (1954) 13, fig. B0; Le6n (1960) 323, fig. in 321. 
Type.—Pacific coast, Colombia, Ouatrecasas 21291 (fruiting speci- 
men), Patino 24 (flowering specimen, para type), 
Small or medium-sized tree up to 15 m. high; growth pseudoapical; 
trunk up to 20 cm. in diameter; primary branches ternate; leafy 
branches ochraceous or ochraceous-ferruginous, or brownish, minutely 
and appressed tomentose, the older glabrate, dark grayish, rather 
smooth or granulate-lenticellate, nitidous, the hornotinous greenish 
ferruginous, tomentose; stipules subcoriaceous, oblong, obtuse, 
striolate, tomentose, about 8-11 mm. long, 2-3 mm. broad. 
Leaves coriaceous, moderately rigid; petiole 9-12 mm. long, thick, 
sub terete, densely and appressed ferruginous-tomentose; blades el- 
liptic-oblong or obovate-ellip tic-oblong, slightly attenuate and asym- 
metrical, or equilateral, rounded at base, rounded or obtuse and 
abruptly acuminate at apex, entire or upwardly sinuate or coarsely 
dentate, 10-32 cm. long, 3-12 cm. broad, the acute acumen 1.5-2.5 
cm. long, above green, pale brown when dry, rather shining, glabrous, 
the costa depressed, thin, the other nerves rather inconspicuous, 
cinereous or greenish cinereous beneath, or pale tawny when dry, the 
costa very prominent, the secondary nerves about 8 on each side very 
CUATRECASAS—CACAO AND ITS ALLIES 573 
Figure 42.—Leaves of Tfuobroma, X%: A, sinuosum (Pavon, s.n.); b, mammosum (Leon 
1363); c, nrmorale (Cuatr. 21291). 
prominent, subparallel, ascending, curving and anastomosing near 
the margin, the transverse tertiary nerves prominent, parallel, the 
nerves of the fourth rank prominulous, broadly reticulate, the minor 
reticulate veins concealed, the whole nervation tomentose, greenish 
ochraceous or subferruginous, by rather thick, tawny, densely dis- 
tributed stellate hairs, the surface within the veins covered by a dense 
and appressed cinereous tomentum of white, fine, minute stellate hairs. 
Inflorescences axillary or on exfoliated branchlets very small, re- 
duced to 1-3 (-5) flowers, the cyme-axis extremely short, the peduncles 
5-10 mm. long, erect, ferruginous-tomentose, 3-bracteolate at apex, the 
pedicels thicker, ochraceous-ferruginous tomentose, 2-5 mm. long, the 
bracts ovate, obtuse, 2.5-3 mm. long, 1.5-2 mm. wide; bracteoles 3, 
broadly cochlear, embracing the bud, orbicular or ovate-rounded, 
574 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
ochraceous-greenish tomentose, 7-11 mm. long and broad, soon 
caducous. 
Sepals thick, ovate-oblong, subacute or subobtuse, 9-10 mm. long, 
4-5 mm. broad, united in the lower third or fourth, first curved- 
patulous, later reflexed, inside purplish or rose, shining, glabrous 
except for the minutely white-tomentose margin, and for the minute 
oblong-capitate, glandular hairs at the base, thick-tomentose outside, 
ochraceous, the stellate hairs with long, thin rays. 
Petal-hoods thick-membranaceous, ochraceous or orange, obovate 
with rounded-cucullate apex, 7-nerved, glabrous and rugose outside, 
minutely hispidulous-pubescent, inside with 7 prominent veins, 3-3.5 
mm. long, 2-3 mm. broad; petal-lamina pedicellate, carnose, rigid, 
red or crimson becoming brownish red, oblong-obovate-spatulate, 
rounded or subtruncate-emarginate at apex, with very minute, sparse 
hairs on the surface and copious longer, slender, weak, spreading hairs 
on the margins, 5-7 mm. long, 3-5 mm. broad; pedicel narrowly 
linear, folded, puberulous, at the broader base barbate, 3 mm. long; 
androecium tube 1.5-2 mm. long; staminodes laminar, rather thick, 
obovate-oblong, rounded or subspatulate at apex, purplish red or 
crimson, covered with sparse, thin, minute hairs, 6-7.5 mm. long, 
3.5-4.5 mm. broad, reflexed in bud, spreading in anthesis; filaments 
moderately thick, 1-1.2 mm. long, pilose towards the base, 3-furcate 
(the branchlets 0.4 mm. long), 3-antheriferous, the anther lobes 
ellipsoid 0.5-0.6 mm, long, connivent; ovary 1.3-1.5 mm. long, oblong- 
ovoid, sub truncate, 5-ridged, thickly ochraceous, hirsute-tomentose; 
styles linear, glabrous, 1.5 mm. long, coherent. 
Fruit ellipsoid, smooth, rounded at apex, slightly constricted above 
the base, 8-10 cm. long, 4.5-6 cm. broad; pericarp coriaceous, rigid, 
1-1.5 mm. thick, when mature fragile, its surface densely appressed 
stellate-tomentose, greenish or bluish, at maturity yellowish, brown 
or tawny; seeds compressed ovoid or subellipsoid, 17-19 mm, long, 
9-12 mm. broad, the surrounding pulp white, yellowish when ripe; 
cotyledons white; fruiting peduncle robust, 1.5-2 cm. long, about 0.8 
mm, thick; germination hypogeous. 
Common names.—"Bacao de monte," "chocolate de monte," 
"bacao," "bacalto," "cacao de monte," "cacaito de monte." 
Uses.—It is said to produce a fairly good chocolate; not known to 
be used by the natives. 
Distribution.—Restricted to the Colombian Pacific coast and 
the Choc6 area between the parallels 3° and 5°3' N. latitude. It is 
recorded from the Calima, San Juan, and Cajambro Rivers. 
This very interesting species is closely related to T. svMncanum and 
T. hylaeum, but it differs specially by its tliree broad, orbicular 
bracteoles subtending the flowers, which before anthesis embrace 
CUATRECASAS—CACAO AND ITS ALLIES 575 
and cover the short-pedicellate bud. This feature is unique in the 
genus Theobroma. 
The leaves of T. nemorale, as well as those of T. hylaeum, are very 
similar to those of T. angustifolium, but they are broader, more oblong- 
elliptic, with a tendency to an obovate shape, and the ochraceous or 
ferruginous hairs beneath are smaller, with shorter, somewhat 
thicker rays than in T. angustifolium. 
COLOMBIA: Choc6: Rfo San Juan, Falestina; part of branch brought in by 
an Indian, one immature fruit 7 x 4.5 cm., 4 VIII 1953, Holliday T/149 (TRIN, 
US). Ibidem, young tree 3 m., sterile, 2 VII 1953, Holliday T/147 (TRIN, 
US). Ibidem, sucker 1J4 m. from leafless fallen trunk, 2 VII1953, Holliday T 148 
(TRIN). Ibidem, Quebrada de las Sierpea; tree 10 m. tall, leaves green yellowish 
above, ashy beneath, 24 IX 1961, Cuatrecasas & Willard 26051 (COL, US). 
Istmina, tree about 7 m., sterile, flowers apparently borne only on branches, 2 
VIII 1953, Holliday & Bartley T/173 (TRIN, US). 
El Valle: Pacific Coast, Rfo Calima, La Trojita, 5-50 m. alt.; small tree; 
leaves coriaceous yellowish green, fruit ovoid-ellipsoid, 8 x 4.5 cm., light brown, 
"bacao de monte," 28 II 1944, Cuatrecasas 16544 (F, VALLE). Rfo Calima, 
Quebrada de la Brea, 20-40 m. alt.; tree 8 m„ leaves subcoriaceous, medium 
green above, green cinereous beneath, nerves green ochraceous, fruits 10 x 6 cm., 
ellipsoid, rounded at apex, constricted at base, tawny, on the branchlets, "choco- 
late de monte," 24 V 1946, Cuatrecasas 21291 (holotype, F; isotypes, B, F, 
VALLE). Ibidem, La Brea, flowering specimens, Patino 24 (paratype, F; type 
of flowers; isoparatypes, F, US). Estaci6n Agroforestal del Calima, 30-50 m. 
alt.; erect tree with abundant handing pods, 8 I 1953, Patifto 117 (US, Herb. 
Cuatr.). Ibidem; small tree 8-10 m., "cacao," "cacafto de monte," "bacaito," 
8 I 1953, PatiOo 116 (US, Herb. Cuatr.). Ibidem; tree with primary ternate 
branches from near the ground; leaves light green; twigs tomentose ochraceous 
or ferruginous, sepals thick, rose or purplish rose inside, ochraceous outside, 
petal-laminae and staminodes thick, rigid, purplish red or dark brown red, hoods 
ochraceous with 7 red veins, "bacao," 23 IX 61, Cuatrecasas Willard 26007 
(COL, US). Cafk> La Brea; young tree 1.5 m., sterile, 29 VI 1953, Holliday 
T/141 (TRIN, US). Estacifin Agroforestal; tree 8-10 m., in land cleared from 
forest, jorquette symmetrical, crimson and yellow flowers borne singly or in pairs 
on small branches, pods 8.5-10 x 5-5.5 cm., fruit peduncle 2 cm. long, 0.75 cm. 
thick, 29 VI 1953, Holliday T/146 (TRIN, US). Pacific coast, Rio Cajambre, 
Silva, Loma de la Vigia, 5-80 m. alt.; small tree, leaves green above, gray beneath, 
"bacao de monte," II-V 1944, Cuatrecasas 17503 (F, VALLE), Ibidem, Que- 
brada del Corosal 0-5 m. alt.; tree 15 m. tall, trunk 20 cm. diameter, leaves 
coriaceous, green above, ashy beneath, fruits ellipsoid rounded at apex, con- 
tracted above the base, smooth, brownish, 10 x 5.5 cm., "chocolate de monte," 
17 V 1944, Cuatrecasas 17738 (F, VALLE). 
TRINIDAD (cult.): Imperial College of Tropical Agriculture, River State 
Diego Martinez, Field 19; 7-8 years old tree, 31 VIII1961, Cuatrecasas, Cope, <fc 
Bartley 25782 T (US). 
20. Theobroma sinuostim Pav<5n ex Huber Figures 41, 42; Map 9; Plate 12 
Theobroma ginuosum Pav6n ex Huber, Bull. Herb. Boiss., II, 6:274. 1906. 
Theobroma 'tessmannii Mildbr. Notizbl. Bot. Card. Berlin 11:139. 1931. 
Theobroma sinuata Ruiz & Pav6n, Fl. Peruv. et Chil. Fol. E, Plate 417, ined. 
576 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Types.—Chicoplaya, Peru, Ruiz <& Pavdn (lectotype, G). Marafidn 
River, Peru, Tessmann 4928 (type of T, tessmannii). 
Large tree with erect trunk; leafy terminal branchlets brownish 
ochraceous when dry, densely hirsute-tomentose, covered by minute, 
whitish, stellate hairs with intricate, slender rays (0.1-0.2 mm. long) 
and by larger stellate, ferruginous hairs with straight, acute, long 
rays (1 mm. long); stipules lanceolate, about 1 cm. long, deciduous. 
Leaves subcoriaceous, firm; petiole short, thick, densely tomentose- 
hirsute, about 1 cm. long; blades obovate-oblong, slightly attenuate, 
rounded and slightly asymmetrical and emarginate at base, sub- 
rounded and abruptly triangular-cuspidate at apex, sinuate-dentate 
in upper third, otherwise entire, about 30 cm. long, 13 cm. broad, 
pale green brownish above, when adult glabrous with abundant 
reddish, punctiforra scars, the midrib and secondary nerves filiform 
impressed, the other nerves less visible, softly tomentose beneath, 
subochraceous, covered by a dense, cinereous layer of minute, white, 
fine, stellate hairs and, especially on nerves, by larger ochraceous, 
stellate hairs, with 4-6, erect, acute, rays 1-1.5 mm. long; the midrib 
thick, prominent, densely tomentose-hirsute, the secondary nerves 
about 9 on each side, ascending, prominent, vanishing near the margin, 
hirsute, the tertiary, transverse nerves prominent, parallel, 5-10 mm. 
distant from each other, the minor veins reticulate and prominulous. 
Sepals ovate-lanceolate united at base, reflexed; petal-hoods obovate, 
concave; petal-lamina obcordate-"trigonous"; staminodes lanceolate- 
obovate; stamens five, 3-antheriferous; ovary hirsute; fruit sub- 
rounded-pyriform, the epicarp smooth, hard, woody, ferruginous 
tomentose. 
The preserved original specimens from the Ruiz and Pavdn herbar- 
ium surely collected by Tafalla have only leaves; they have been used 
by me for the above description. The short description of flowering 
characters has been taken from the manuscript of the unpublished 
Flora of Peru and Chile of Ruiz and Pavdn. This description was 
written by Ruiz or Pavdn based on data sent by Tafalla. Some of 
the data were misinterpreted by the authors who describe the anthers 
"quinque in singulo filamento," and this is not the case. Tafalla 
wrote in his "notas" that each filament was divided in six "lacinias" 
bearing one anther each, and that the number of stamens was five in 
T. alba, sinvata, and cordata, whereas it was ten in T. digifMa. Pavdn 
described the inflorescences as being cauline; we have no basis either 
to affirm or deny that assertion. The petal-lamina is described as 
"trigona"; probably it was slightly 3-dentate, and this feature was 
extremely exaggerated by Pulgar in his drawing. 
Theobroma sinuosum, because of the little information available, was 
disregarded by most of the authors who listed cacao species; Chevalier 
CUATRECASAS—CACAO AND ITS ALLIES 577 
who saw specimens of it considered it as a synonym of T. ferrugineum, 
which was not well defined by him in his revision. My study of the 
sterile specimens of the type convinces me that T. sinuosum is a very 
different species, unique in possessing an indument comparable to 
that of the young plants of T. chocoense and T. simiarum. On the 
other hand, after careful study of the description and photograph of 
T. tessmannii Mildbr. which I had formerly associated with T. subin- 
canum, I arrived at the conclusion that the Tessmann specimen is 
definitely distinct from T. subincanum and that it coincides with T. 
sinuosum. Unfortunately the Tessmann specimens are not existing 
any more, but the Mildbraed's detailed description of the indument 
permits us to differentiate his species from the closest allied species 
growing in the same area, namely T. subincanum. Some doubts 
may remain about the identity of T. sinuosum and T. tessmannii. 
The Tessmann plant definitely had axillary inflorescences, whereas 
the Pav6n plant was described as being cauliflorous, but the species 
could well have both cauline and axillary inflorescences. The geo- 
graphical range and the identical kind of indument are the reasons 
that I consider the species synonymous. 
I wish to supplement the description given above with the data 
taken from Mildbraed's description: Leaves subcordate at base, 
25-35 cm. long, 8-12 cm. broad, with about 10 secondary nerves; 
sepals lanceolate, acute, about 10 x 4 mm., connate at base for 2 mm.; 
petal-hood whitish, lamina red, subquadrate-rounded and apiculate, 
3 x 3 or 2 x 3 mm.; staminodes dark red, subspatulate-elliptic, 6 mm. 
long, 3 mm. broad; stamens 3-antheriferous; pedicels 2 cm. long. 
Common names.—Cacao de monte, Pako Kakao (Tessmann). 
Uses: The pulp of the fruits is eaten by the natives. 
Distribution.—Upper river valleys of the Huallaga and Marafi6n 
in Peru, 
PERU: Chicoplaya, "Pavdn," collected by Tafalla (lectotype, G; BM). Rio 
Marafi6n from Iquitos to the mouth of Santiago, near Pongo de Mansariche, 
ca. 77*30' W,, Tessmann 4928 (type of T, tessmannii, photo F. M. 17942). 
21. Thcobroma canumanense Pires et Fr6es, sp. nov. Figure 43; Map 9 
Arbor 18 m. alta pauciramosa ramulis dense ferrugineo-tomentosis, 
pilis crassiusculis mediocribus stellatis 8-14 radiis acutis 0.3-0.7 mm. 
longis instructis. 
Folia rigide coriacea, petiolo crasso dense ferrugineo-tomentoso 
8-10 mm. longo. Lamina oblongo-obovata vel obovato-elliptica basi 
paulo angustata rotundataque leviter asymmetrica apice subrotundata 
subite breviterque acuminata margine Integra vel sursum leviter 
grosseque dentata, 8-20 cm. Ionga, 3-8.5 cm. lata, supra in sicco 
tabacina subnitida leviter rugosa juventute pilosa deinde glaberrima, 
costa nervisque secundariis filiformibus impresses ceteris paulo con- 
578 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
spicuis, subtus ferrugineo-tomentosa, costa valde elevata, nervis 
secundariis 5 vel 6 utroque latere elevatissimis duobus basilaribus 
ascendentibus ceteris patulo-ascendentibus arcuatisque ad marginem 
decurrentibus anastomosantibus, nervis tertiariis transversis bene 
elevatis 3-6 mm. inter se distantibus, nervulis quaternariis elevatis 
transversis, minoribus prominenter minuteque reticulatis, pilis dimor- 
phis: a) pilis stellatis albidis minutissimis dense intricatis areolas 
tectis, b) pilis ferrugineis crassiusculis mediocribus stellatis radiis 
patulis supra nervationem copiosissimis. 
Flores in cymis axillaribus vel extra-axillaribus paucifloris congeste 
glomerati; pedunculi ad 6 mm. longi tomentosi apice 3-bracteolati, 
bracteolis 1-1.5 mm. longis triangularibus tomentosis; pedicelli 
tomentosi 1 mm. longi. Al&bastra globosa crasse ferrugineo-tomentosa 
circa 5 mm. diam. Calyx sepalis crassiusculis carnosis in anthesin 
subpatulis paulo reflexis, circa 7 mm. longis, 4 mm. latis, basi 1.5 mm. 
longitudinaliter coalitis, intus glabris basi excepto, pilis densis crassis 
glandulosis ad marginem insertion© praeditis extus dense stellato- 
tomentosis margine dense minutissime pilosulis. 
Cucullus petali obovoideo-ellipsoideus circa 3-3.5 mm. longus, 
2-2.2 mm. latus, carnosulus extus glaber intus 7 costis elevatis stri- 
gulosis instructus. Lamina petaloidea rubra crassa semirotundata 
apice emarginata, circa 2 mm. longa, 3 mm. lata, sursum utrinque 
minutissime pilis margine ciliata, basi subite in pediculum circa 2 mm. 
Longum longe ciliatum contracta. 
Androecium rubrum; tubus circa 1.5-1.6 mm. altus glaber. Sta- 
minodia crassa cttrvata subspathulata sursum dilatata apice leviter 
emarginata, circa 6 mm. longa, 2.4-2.8 mm. lata, margine longe 
flexuoso-ciliata cetera glabra. Stamina filamentis glabris crassis, 
1.2-1.4 mm. longis triantheriferis, lobis antherae ellipsoideis circa 
0.4 mm. longis. Ovarium 1.2-1.5 mm. longum obovatum pentagonum 
in angulis et subapicem hirsutulum, Styli circa 2 mm. longi glabri 
versus apicem plus minusve liberi. Fructus ignotus. 
Type in the U.S. National Herbarium, No. 2404642, collected in low, 
firm land, Rio Canumao, tributary of Madeira River, municipality of 
Borba, State of Amazonas, Brazil, October 5, 1957, by R. L. Frdes 
(No. 33783). Isotype in the herbarium of Instituto Agronomico 
do Norte, Belem do Par6,. 
Theobroma canumanense is closely related to T, sinuosum; its 
leaves and indumentum conform well with those of the type of this 
species showing only some differences due to the fact that the T. 
sinuosum specimens came from a young plant. The vegetative 
characters of the Fr<5es plant also coincide with the description of 
T. tessmannii given by Mildbraed, but I found a few differences 
which indicate that the Fr6es specimens belong to a different species. 
CUATRECASAS—CACAO AND ITS ALLIES 579 
Figure 43.—Theobroma canumanense (Froes 33783): a, petal, inside view; b, petal, lateral 
view, X 5; c, androecium, X 5; D, stamen, X 10; e, sepals, inside view, X 2; f, pistil, 
X 5; g, bud, X 2. 
T. canumanense has smaller flowers, the sepals being only 7 mm. 
long; they are 10 mm. in T. tessmannii which also has a longer 
androecium (8.5 mm. high); the petals and staminodes are glabrous 
in the Tessmann plant according to the Mildbraed description and 
drawings, while they are long-ciliate in T. canumanense. The 
inflorescences are more compact in this species than in T. tessmannii. 
The photograph of the latter shows the adult leaves to be subcordate 
and broader at base and provided with one or two additional pairs of 
basal veins (not conspicuous in T. canumanense); the secondary 
nerves are more numerous (8-11 pairs) in Tessmann's plants. 
T. canumanense also differs from all related species by its extremely 
short pedicels (up to 1 mm. long). 
Common names.-—Names and uses not recorded. 
Distribution.—Limited to the region of Rio CanumSo, a tribu- 
tary of the Bio Madeira in Brazil. 
BRAZIL: Amazonas: Regi&o do Rio Madeira, Rio CanumSo, munieipio de 
Borba, 5 XI1957, R. L. Frdes 33783 (US, holotype; IAN, isotype). 
Section 6. Andropetalum 
Theobroma sect. Andropetalum Cuatr., sect, no v. Figure 4 
Lamina petalorum anguste spathulata longe attenuato-stipitata; 
cucullum 7-nervatum; staminodia crassa latissima petaloidea arcuato- 
reflexa petala obtegentia; stamina 3-antherifera; fructus ellipsoideo- 
oblongus laevis tomentosus supra basim constrictus ad apicem 
angustato-mammillatus epicarpio duro lignoso; semina germinatione 
hypogaea; folia subtus adpresse stellato-tomentosa; infiorescentiae 
axillares brevissimae; caulis incrementum pseudoapicale ,* rami 
580 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
primarii ternati; calyx cymbiformis 3-lobatus, sepalis usque ad 
tertiam partem Tel ad medium connatis. 
Type species.—Theobroma rnammosum Cuatr. & Le6n. 
This section comprises a single species. It is characterized by the 
broad, obovate-spatulate staminodes (as broad as long), which are 
reflexed even during an thesis, completely covering the petals, and 
by the reduction of the petal-laminae, which are narrower and 
smaller than in the closely related section Glossopetalum. 
22. Theobroma rnammosum Cuatr. & Le6n 
Figures 6, 36, 41, 42, 44; Map 9 
Theobroma rnammosum Cuatr. & Le6n, in Le6n, I list. Interamer. Cienc. 
Agr. Bol. TAcn. 2:1-6, figs. 1949; Le6n (1960) in 320, 317, Jig, 
Type.—Siquerres, Lim6n, Costa Kica, Ledn 291. 
Small tree, 6-7 m. high; trunk about 25 cm. in diameter with 
rather smooth, brown bark 1 mm. thick and white, hard wood; growth 
pseudoapical; branches ternate, from near the base, spreading or 
more or less descending; terminal branchlets grayish or somewhat 
ochraceous, appressed and minutely stellate-tomentose, later gla- 
brous, gray, rugose; stipules subulate, acute, 4-5 (-10) mm. long, 0.6 
mm. wide, deciduous. 
Leaves subcoriaceous; petiole rather thick, subterete, straight or 
somewhat flexuous, ochraceous or ferruginous, stellate-tomentose, 5- 
12 mm. long; blade elliptic-oblong or oblanceolate, slightly attenuate 
to the apex and suddenly acuminate, slightly narrowed toward the 
asymmetrical base, rounded in one side, subrounded or subcuneate 
at the other side, the margin entire or sinuate-dentate near the apex, 
10-25 cm. long, 3.5-10.5 cm. broad, including the acumen, this acute, 
8-20 mm. long, glabrous above, dark green, when dry pale brownish, 
the costa filiform, depressed, the other nerves hardly noticeable, 
cinereous beneath, except for the main nerves densely and appressed 
stellate-tomentose, covered by thin and minute, white stellate hairs, 
the costa very prominent, the prominent secondary nerves 9-12 on 
each side, regularly parallel, subascending, curved and vanishing near 
the margin, the transverse tertiary nerves filiform, prominent, the 
lesser veins reticulate, thin, prominulous, but covered by the tomen- 
tum, the midrib, secondary, and tertiary nerves with scattered or 
copious, larger, thicker, spreading, stellate hairs. 
Inflorescences very small, axillary, cymose with few (usually 2) 
flowers (1-3), the axis extremely short, tuberculate, giving rise usually 
to a single ferruginous-tomentose branch 8-12 mm. long, this 2- or 3- 
furcate at apex into 2 or 3 peduncles; peduncles very short, about 1 
mm. long, 3-bracteolate at apex, each articulate to a pedicel; bracteoles 
linear, 2-4 mm. long, tomentose; pedicels rather thick, tomentose, 
CUATRECASAS—CACAO AND ITS ALLIES 581 
Figure 44.—Thtcbroma mammosum (Leon 1363): a, petal, inside view, X 5; b, petal, 
laterally, X 5; c, part of androectum, X 5; D, flower in antbesis, X 2.5; e, gynoecium, 
X 5. 
8-12 mm. long; buds globose, densely ferruginous tomentose, about 
1 cm. in diameter. 
Sepals 11-12 mm. long, 5-6 mm. broad, thick, triangular-ovate, 
united completely by pairs or by 2 and 3, and all five in the lower 
third, forming a cupular, umbilicate, 2 or 3 short-lobate calyx, yellow- 
ish green inside, red tinged, glabrous except for the glandular, thick, 
erect, fasciculate, congested trichomes at the inner base, these 0.4- 
0.7 mm. long, densely, thickly, stellate-tomentose outside, greenish 
ochraceous or ferruginous. 
Petal-hoods about 7-8 mm. long, 5-7 mm. broad, elliptic-obovate, 
broadly rounded-cucullate at apex, dark red, rather carnose with 
7 thick nerves, between the nerves veined with yellowish or rose by 
transmitted light, minutely papillose inside, rugulose outside with 
sparse, long, acute, straight or slightly flexuous hairs throughout, 
these more copious at margin; petal-lamina erect in full an thesis, 
narrow, thick, dark red or purplish red, truncate-spatulate or slightly 
emarginate, usually plicate, with long, ferruginous, slightly flexuous 
hairs, these scattered outside, abundant at margin and inside, 3 mm. 
long, 2.5-3 mm. broad, gradually narrowed toward the base into a 
plicate pedicel about 4 mm. long, 0.5 mm. wide. 
582 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Androecium purplish red or dark red, the tube carnose, about 
4 mm. high, 3-4 mm, broad, glabrous; staminodes large, petaloid, 
reflexed, the lower part thick, carnose, toward the apex gradually 
thinner, membranaceous, transparent-veined, obovate-spatulate, sub- 
truncate and slightly sinuate at apex, laterally usually 1-dentate, the 
lower half suddenly narrower, about 11-12 mm. long, 10-12 mm. 
broad at top and 2.5 mm. wide at base, glabrous or with sparse hairs 
near the top outside, with copious, ferruginous, simple or stellate, 
rather long hairs inside; filaments thick, glabrous, about 1 mm. long, 
shortly 3-furcate, 3-antheriferous, the lower branchlet extremely 
short, the other two lateral about 1 mm. long; anther lobes ellipsoid, 
suborbicular, about 1 mm. long after an thesis; ovary subglobose, 
slightly 5-angulate, whitish, densely tomentose-hirsute, about 1.8 
mm. high and thick; styles 2 mm. long, glabrous, erect, acute, con- 
nivent, only united at base. 
Fruit at maturity 16-22 cm. long, 6-8.5 cm. broad, cylindric-oblong, 
terete or seldom slightly pentagonous, broad and umbilicate at base, 
strongly contracted and slightly pentagonal above the base, suddenly 
narrowed near the apex forming a mammiform, umbilicate end 2-3 
cm. long and 2-2.5 cm. broad. Pericarp coriaceous, hard, smooth 
or slightly verrucose downward, pale green, covered by dense, short, 
ferruginous tomentum, the woody epicarp 1.5-2 mm. thick with a 
tomentose epiderm outside, the mesoendocarp carnose, white, about 
7 mm. thick; pulp enveloping each seed fleshy, fibrous, white; seeds 
ovoid-amygdaloid, brownish, 22-27 mm. long, 14-17 mm. broad, 
10-13 mm. thick; cotyledons white; germination hypogeous. 
Common name.—Cacao silvestre. 
Distribution.—Eastern coastal mountains of Costa Rica, where 
it is extremely rare in primary forest. It has been found wild only 
twice, at altitudes from 300 to 800 meters. Cultivated in few agri- 
cultural experimental stations, 
COSTA RICA: Lim6n: La Lola, 100 m. alt., 12 V 1948, Escamtlla, s.n. (MO). 
Sfquirres, Finca La Lola, 300 m. alt., I 1949, Le6n 291 (holotype, TURRI). 
La Lola, cerca de Madre de Dios, 100 m. alt., cultivated, 30 I 1949,Le6n 1363 
(paratype, F, TURRI). Ibidem, Experimental Station I.I.C.A., cultivated; 
tree 6 m., stem 20-25 cm. diam., leaves chartaceous, firm, green above, pale 
beneath, branches ternate, growth pseudoapical, flowers dark red and brown red, 
fruit ellipsoid, contracted at apex, 6 XI 1961, Cuairecasas tfc Paredes 26535 (US). 
Heredia: Puerto Viejo, Sarapiquf, 700-800 m. alt., IV 1959, Holdridge 
146 (TURRI). 
Cartago: Tunrialba, grounds of I.I.C.A., cultivated, 600 m. alt., 4 XI 1961, 
Cnatrecasas & Le6n 26516 (US). 
TRINIDAD: River State Diego Martinez, I.C.T.A., Field 19, cultivated from 
seed received from Bel6m do Par£; tree 8 m. high, stem rugose-tuberculate, brown, 
abundant dry fruits hanging from branchlets, 1 IX 1961, Cuotrecoaoa & Cope 
25791 (US). 
CUATRECASAS—CACAO AND ITS ALLIES 583 
Hybrids 
Theobroma angustUolium Moc. & Sess6 cf X mammosum Cuatr* & Le6n 
Trinidad, I.C.T.A., Cuatrecasas <& Cope 25800. Well-developed 
tree, with intermediate characters. 
Theobroma grandiflorum (Spreng.) Schum. X obovatum Klotzsch. 
Bel&n do Par6, Brazil, Museu Goeldi, arvore 480, 22 XII 1958, 
Cavalcante 937 (MG, US). 
Theobroma grandiflorum (Spreng.) Schum. 9 X subincanum Mart. cP. 
Hybrid fertile obtained by Addison and Miranda (1951) 13. In- 
termediate features of parents. 
Theobroma grandiflorum (Spreng.) Schum. 9 X obovatum Kl. cf. 
Brazil. Fertile hybrid obtained by Addison (1951) 10, pi. Sf 
pi. 2, jig. C. F. cultivated at Bel6m do Par6, 6 XI 52, Pires 4343 
(IAN). Ibidem, Pires 4344 (IAN). 
Theobroma mammosum Cuatr. & Le<5n cT X simiarum Donn. Smith.? 
Costa Rica, Turrialba IICA, robust young trees obtained by Soria. 
Theobroma mammosum Cuatr. & Le6n 9 X simiarum Donn. Smith c", 
Costa Rica, Turrialba, IICA, robust young trees obtained by Soria. 
Theobroma obovatum Klotzsch d* X subincanum Mart. 9. 
Fertile hybrid obtained by Addison and Miranda (1951) 14, pi. IS, 
jig. 13, pi. 4t M' 0. Also found spontaneous by Cope & Holliday 
in Colombia: Rio Caquetd, Remolino, tree ll-12m., with some 
characters of T. obovatum and others of T. subincanumt 2 V 1953; 
it has almost smooth pods but bears flocose, woolly pubescence on 
the young shoots and leaves, Holliday dk Cope T122 (COL, TRIN, US). 
Caquetd, Rio Cagudn, tree 8 m., 27 IV 1953, Cope <& Holliday T 117 
(COL, TRIN). Brazil, Amazonas, Fonte BOa; tree 15 m. alt. "cu- 
purana," Frdes 20648 (IAN, US). Trinidad, ICTA, experimental 
camps, Cuatrecasas cfc Cope 25797 (US). 
Theobroma speciosum Willd. ex Spreng. X sylvestre Mart. 9* 
Bel6m do Par6, Brazil, 6 XI 1952, Pires 4345 (IAN, COL). Ex- 
perimental hybrid by Addison. It has intermediate characters as 
described and illustrated by Addison & Miranda (1951) 14, pi. 6 
fiQ' pl* ^ 
Theobroma speciosum Willd. ex Spreng. 9 X sylvestre Mart. 
Beldm do Par6, Brazil. Experimental hybrid obtained by Addison, 
similar to the former. Addison & Miranda (1951) 15. 
At the Institute Interamericano, Costa Rica, Dr. Soria (1961) 
tried to hybridize T. cacao with T. angustifolium, mammosum, simia- 
rum, and bicolor. The cross T. angustifolium X cacao $ gave small 
680-695—64 14 
584 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
plants which did not grow more than about 10 cm. in two years. 
Theobroma cacao 9 X mammosum <? gave fruits, but the seeds were 
very weak; the same happened with T. cacao $ X simiarum d\ The 
cross T. bicolor cT X cacao 9 was negative. See also Addison and 
Miranda in the Historical Sketch above. 
Rejected Names and Excluded Species 
Theobroma Kalagua de Wildeman, Bull. Herb. Boiss. 7:957, pi. 11. 1899. 
Type.—Panama, Patin, s.n. (lectotype, BR, leaf) (F. M. Photo 
40742). 
This species was described by de Wildeman as very remarkable, 
because of the extraordinary combination of flowers similar to those of 
T. angustifolium and T. simiarum, fruits resembling T. simiarum, 
and quite different leaves which although resembling those of T. 
angustifolium in shape differed by the lack of pubescence. But 
Patin gave assurance to Wildeman that these different parts came 
from a single tree. De Wildeman wrote: "qui appartiendraient sans 
le moindre doute & la mfime plante. Une confusion aurait cependant 
pu 6tre possible parce que le Tk. simiarum existe dgalement en 
Colombie, od M. Ch. Patin, qui s'adonne a la recherche des plantes 
utiles, Pa d&ouvert k Choco, province de Cauca" (Wild. p. 958). 
In the U.S. National Herbarium, attached to a herbarium sheet 
(1,382,338), is a letter sent to J. Donnell Smith by Ch. Patin, dated 
Brussels 19 Oct. 1900. Here Patin states "As concerns T. Kalagua 
. . . there was a doubt about the leaves used to determine the specy 
[sic]: the young plants which we have just got here from seeds have 
proved that it occurred really a mistake in the description made of 
the leaves brought to me by my collector." This contradicts the 
earlier assurances given by Patin about all his specimens having been 
collected from a single tree and shows that he was not the collector 
himself. Later on Patin writes, "I think that the T. Kalagua is 
just your T. Simiarum " Patin never explained where his specimens 
were collected. Probably they came from the region of PanamA. 
Regarding the specimens sent with the above cited letter, the seedling 
leaves and small fragments of fruits might well be those of X. simiarum. 
To clear up this question, in 1953 I asked Dr. Robyns, director of 
the Jardin Botanique de l'Etat, Brussels, to send me the material 
of T. kalagua preserved at that herbarium. Very kindly, Dr. Robyns 
sent me the only existing herbarium sheet of the type material of 
T. kalagua, which is represented only by one leaf, undoubtedly the 
same used to illustrate the plate in the publication. To a further 
request to Dr. Robyns the answer was given that no other material 
CUATRE CABAS—CACAO AND ITS ALLIES 
585 
of this species existed except for some fruits which "selon toute 
vraisemblance appartiennent au materiel Patin s.n. recolt£ en 1899, 
se trouvaient dans nos collections de fruits." 
Examination of the Patin specimens proves that the leaf mounted 
on this sheet at BR (photo F. M. No. 40742) is the original Patin leaf 
used for de Wilde man's illustration and that this leaf belongs to 
Theobroma cacao. This leaf is the only unquestionable part of the 
type apparently existing, and I select it as the lectotype. The three 
fruits received on loan from Brussels labeled T. kalagua were: 1) A 
half shell, to be discarded as belonging to Theobroma bicolor. 2) An 
entire pod with a modern label from "J. Bot. Br." reading "Th. 
Kalagua Wild. Colombie, sans date Coll. Patin." This fruit is 
smooth, ellipsoid, oblong, slightly attenuate and rounded at apex and 
broadly rounded, umbilicate at base, 18-20 cm. long, and 8.8 cm. 
broad; the surface is tomentose, the woody epicarp about 1.5 mm. 
thick, the dried, spongy mesoendocarp 2-4 mm. thick; a fragment of 
this shell is lacking, and this could be the fragment of fruit sent by 
Patin to Donnell Smith and preserved at the U.S. National Herbarium 
(1,382,338). This fruit is not that used for the illustration, because 
the original was sectioned; furthermore, this pod looks somewhat 
more oblong than the one figured in the plate. It may well belong to 
T. simiarum, as may also the seedling leaf sent to Donnell Smith. 
3) The third fruit received from Brussels bears two labels; one reads 
(t
 Theobroma kalagua, Colombie, Choc 6, Pro v. de Cauca, leg. Ch. 
Patin 1899," and the number 823; the other label reads "Theobroma 
simiarum D. Sm. Colombie, leg. Ch. Patin"; it consists of half a shell 
whose section is ovate-ellipsoid, 19 cm. long and 14.2 cm. broad and 
about 2 cm. thick; the woody epicarp is 1.5-2 mm. thick; the thick 
mesoendocarp is compact. It is different from all species of Theo- 
broma known to date; it might be an undescribed species, but it could 
also belong to another genus. 
In conclusion, we may infer that Patin gathered several specimens 
of different Theobroma species collected by different persons and 
coming from several places (Panama, Chocd . . .). The flowers 
described by de Wildeman could well be flowers of T. angustifolium 
or T. simiarum; the possibility of their growing in Panama, perhaps 
planted, cannot be discarded. They also could have been flowers 
from trees of T. stipulatum, T. chocoense, or from some undescribed 
species. An important character of the flower, its color, was 
not mentioned by Patin. The fruit shown in the plate is thicker 
than that of T. simiarum; it recalls very much that of T. grandijlorum, 
a species more or less widespread in gardens, and it is also similar to 
that of T. chocoense and T. stipulatum. It may also belong to an 
undescribed species. To ascertain to what species such a fruit belongs, 
586 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
it would be necessary to see at least two associated organs from the 
same tree, that is to say fruit and leaves, fruit and flowers, or, also, 
flowers and leaves. The above comments on the available data 
prove the disorderly way of Pa tin's work and his unreliability. 
There is no reason to believe that the flowers and fruits of Patin's 
collections were brought from the same tree or from the same species. 
Patin never mentioned the locality of the specimens sent to de Wilde- 
man or who collected them. Colombia is always cited but at that 
time Panama was part of Colombia. At present the only identifiable 
Patin specimen from his syntypes is the leaf (BR) and that is T. 
cacao. Consequently, T. kalagua becomes a synonym of T. cacao. 
Theobroma guianense (Aubl.) Grnel., Syst. Veg. ed 13, 2:1151. 1796. 
Cacao guianensis Aubl. PI. Guian, 2: 683, pi. 275. 1775. 
Syntypes.—Aublet illustrations (I.e.) and specimens at British 
Museum (Natural History). 
Under Cacao guianensis, Aublet gave a detailed and illustrated 
description made up of a mixture of three different species. There is 
complete agreement between the description of each part and the 
corresponding illustration. The flowers were described from specimens 
of Theobroma cacao L.; the branches and leaves from T. subincanum 
Martius; the fruits cannot be identified with any other species included 
in Bernoulli's and Schumann's treatments. The illustration of this 
fruit (PL 275, jigs. 16 & 17) agrees unmistakably with only one recent 
collection of Theobroma fruits, namely that from French Guiana by 
Benoist, which is the type of T. velutinum Benoiat. Aublet's short 
diagnoses of the fruit, especially the French description "I'ovaire 
devient une capsule ovoide & cinq arretes arrondies saillantes" also 
agree perfectly with it. It seems that Aublet considered the most 
typical part of his "species" the fruit, since he headed the description 
of his Cacao guianensis with a short definition based only on the 
fruit: "CACAO (Guianensis) fructu ovato, quinquangulari, tomen- 
toso, rufescente (Tabula 275)" (p. 683); furthermore, as Sandwith 
pointed out, the French name given by Aublet [Le Cacaoier anguleux. 
(Planche 275)] was taken from the same diagnosis. Thanks to Benoist 
we know now that this kind of fruit belongs to a species with a kind 
of leaves very different from those described by Aublet, the species 
described by Benoist as T. velutinum. 
But the nomenclatural problems have to be solved on the basis of 
types according to the International Code. At the British Museum 
there are preserved Aublet specimens and among them type specimens 
of Cacao guianensis which are syntypes. In 1954, I could identify 
by close examination an herbarium sheet (with foliage, one flower, 
CUATRECASAS—CACAO AND ITS ALLIES 587 
and an immature fruit) as belonging to T. subincanum Mart.; this 
specimen agrees with the description and drawings given by Aublet 
for the leaves and branchlets; this evidence could easily incline 
us to use this specimen as lectotype for 0. guianensis. But in the 
carpological collection of the British Museum there is a fragmentary 
fruit, also a syntype of Aublet's species, which belongs to T. cacao. 
Until now, there has been only confusion about the identity of 
Theobroma guianense. Bernoulli (p. 7) wrote: "Cacao guianensis 
bleibt somit eine vollstaendig ungewisse Art. Sie scheint auch von 
keinem weitern Autor gesehen worden zu sein, sondern immer nach 
Aublet citiert zu werden." Schumann placed it as synonym of 
Theobroma cacao, surely on account of its flowers. Chevalier con- 
sidered the species synonymous with T. speciosum Willd., erroneously 
interpreting Aublet's descriptions and drawings of the leaves and a 
photograph of the Aublet herbarium specimen (syntype) at the 
British Museum; these, as I have pointed out above, belong to T. 
svbincanum Mart. He also identified the Benoist collections of T. 
vdutinum erroneously as T. speciosum. 
Consequently, Cacao guianensis Aubl. is not a true species, but a 
mixture of three species. Therefore the Aublet "species" and name 
has to be rejected as "nomen confusum" (articles 63(3), 65, and 66 
of the Code of Nomenclature). The name Theobroma guianense 
has never been consistently used in monographs and general books 
for any known species. 
"Theobroma fossilium" Berry, Proc. U.S. Nat. Mus. 75(24) :8, pi. 1, jigs. IS, 14• 
1929. 
In regard to this unfortunate name, the following opinion of R. W. 
Brown, former paleobotanist of the U.S. Geological Survey, is final: 
"This specimen, considered by Berry to be the first fossil record of 
Theobroma, is a section of the forepart of a reptilian jaw. Berry 
mistook the bony structure for the pulp, and the teeth for the seeds 
of a chocolate-bean pod. Although described among Tertiary 
plants, the specimen, as stated by Berry, came from near Leiva, 
Department of Boyacd, Colombia, where Cretaceous strata crop out." 
(Journ. Washington A cad. Sci. 36:353. 1946). 
Theobroma alba Bernoulli, Uebera. Art. Theobroma 14. 1869; Jumelle (1899) 
35; De Wildem&n (1902) 98. 1902. 
Type.—Essequebo et Cuyaunic, British Guiana, C. F. Appun 1, 
1860 (holotype, Herbarium Hookerianum, K). 
The type of this species consists of leaves of a young, sterile plant 
of the genus Licania. I have identified it by comparison from the 
abundant material existing in the Royal Botanic Gardens, Kew, with 
588 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
L. venosa Rusby, a widespread species of the Guianas. The following 
new combination is necessary: 
Licania alba (Bernoulli) Cuatr., comb, no v. Theobroma alba Bernoulli, Uebers. 
Art. Theobroma 14. 1869. Licania venosa Rugby, Descr. New Sp. So. 
Am. Pi. £6, 1920. 
Theobroma albiflorum (Goudot) De Wildeman, PI. Trop. Gr. Cult. 90. 1902= 
Herrania albifiora Goudot, Ann. Sci. Nat. Ill, 2:230, pi. 5. 1844. 
Theobroma aspera (Karsten et Triana ex Triana) Van Hall, Cacao, ed. 2, p. 49. 
1932. (Brotobroma aspera Karsten et Triana ex Triana, Nuev. jen. y esp. 
fl. Neo-Granat. 12. 1854) = Herrania nitida (Poepp.) Schultes, Caldasia 
2:16, 17, pi 1943. 
Theobroma augusta L. Syst. Nat. 3:233. 1776; Willd. Sp. PI. 3:1424. 1803— 
Abroma augusta (L.) L. f. Suppl, 341. 1781. 
Theobroma balaensis (Preuss) De Wildeman, PI. Trop. Gr. Cult. 89. 1902= 
Herrania balaensis Preuss, Exped. Centr. und SGd-Amerika 253, pi. 7. 
1901. 
Theobroma camargoanum (Schultes) Ducke, Bol. T6cn. Inst. Agron. Norte 
28:15. 1954=Herrania camargoana Schultes, Bot. Mus. Leafl. Harvard 
Univ. 14:120, pi. 29, SB. 1950. 
Theobroma eeltifolia Salisburg, Pro dr. 387. 1796=6rt«izuma ulmifolia Lam. 
Encycl. M6th. 3:52. 1789. 
Theobroma guazuma L. Sp. PI. 782. 1753=Guazuma tomentosa H. B. K. Nov. 
Gen. Sp, 5:320. 1823. Freytag (1951) 214. 
Theobroma hastata Chevalier, Rev. Bot. Appl, 26:273. 1946, nomen nudum; 
lapsus calami for T. sagittala Pav6n ex Chevalier. 
Theobroma laciniifolium (Goudot ex Triana et Planchon) De Wildeman, PI. 
Trop. Gr. Cult. 90. 1902=Herrania lactniifolia Goudot ex Triana et 
Planchon, Pro dr. Fl. N. Granat. 209. 1862. 
Theobroma mariae (Martius) Schumann in Mart. Fl. Bras. 12?:71, pi. IS. 
1886—Herrania mariae (Mart.) Decaisne ex Goudot Ann. Sci. Nat. Ill, 
2:233. 1844. 
Theobroma montana Goudot ex Bernoulli, Uebers. Art. Theobroma 15. 1869, 
nomen nudum. No description. 
Theobroma nitidum (Poepp. et End!.) Schumann in Mart. Fl. Bras. 123:72. 
1886. (Abroma nitida Poepp. et Endl., Nov. Gen. Sp. PI. 3:73. 1845) = 
Herrania nitida (Poepp. et Endl.) Schultes, Caldasia 2:16,17, pi. 1943. 
Theobroma purpureum Pittier, Repert. Sp. Nov. Fedde 13:319, 1914= 
Herrania purpurea(Pittier) Schultes, Caldasia 2:333. 1944. 
Theobroma pulcherrimum (Goudot) De Wildeman, PI. Trop. Gr. Cult. 89. 
1902=Herrania pulcherrima Goudot, Ann. Sci. Nat. Ill, 2:232, pi 6. 
1844. 
Theobroma saglttata Pav<5n ex Chevalier, Rev. Bot. Appliq. 26:274. 1946, 
nomen nudum=Herrania nitida (Poepp. et Endl.) Schultes, Caldasia 2:16, 
17, pL 1943. 
Theobroma tomentosa (H. B, K.) Gdmez, An. Hist. Nat. 19:217. 1890= 
Ouazuma tomentosa H. B. K. Nov. Gen. Sp. 6:320. 1823. 
Theobroma undulata Pavdn ex Chevalier. Rev. Bot. Appliq, 26:268. 1946, 
nomen nudum: lapsus calami for T. sinuo&um Pavdn ex Huber. 
Collections Cited 
AcoSTA SOLIs, M., & Giler, M. 
12392 gileri 
12423 gileri 
Allen, P. H. 
3105 grandiflorum 
4593 bicolor 
6259 aDgustifolium 
6341 angustifolium 
Appun, C. F. 
1 Alba 
Araqtje, J., & BAEKLEY, F. A. 
18C745 hylaeum 
Archer, W. A. 
7517 subincanum 
7537 obovatum 
7549 grandiflorum 
7551 microcarpum 
7619 BpecioBum 
7721 speciosum 
7734 grandiflorum 
7820 subincanum 
Abplund, E. 
10271 bicolor 
12911 subincanum 
13410 subincanum 
Aublet, J. B. C. F. 
s.n. cacao (fruit) 
e.n. subincanum 
BAFOG (Bureau Aob. et Forestier 
Gutanais) 
136M velutinum 
7386 velutinum 
BAILEY L. H. 
s.n. angustifolium 
Baker, C. F. 
62 grandiflorum 
421 grandiflorum 
2102 angustifolium 
Bakeb, R. E. D. 
16 grandiflorum 
33 subincanum 
34 bicolor 
37 glaucum 
38 subincanum 
s.n. angustifolium 
BAKEB, R. E. D., Baktlet, B. G., & 
Holliday, P. C. 
31 subincanum 
32a subincanum 
BAKEB, R. E. D., & COPE, F, W. 
2 bicolor 
3 subincanum 
4 subincanum 
5 grandiflorum 
6 grandiflorum 
7 subincanum 
11 glaucum 
11a bicolor 
12 subincanum 
13 subincanum 
21 subincanum 
25 subincanum 
26 bicolor 
27 obovatum 
28 microcarpum 
29 microcarpum 
30 obovatum 
31 subincanum 
32 subincanum 
Bartley, B. G., & Holliday, P. C, 
T 46 grandiflorum 
T 47 bicolor 
T 65 subincanum 
T 66 bicolor 
T 68 subincanum 
T 69 glaucum 
T 70 glaucum 
T 71 subincanum 
T 72 subincanum 
T 74 glaucum 
T 75 subincanum 
T 166 hylaeum 
Bbnoist, R. 
516 velutinum 
BERNOULLI, G. 
94 bicolor 
95 angustifolium 
Bernoulli, G., & Cario, R. 
3188 angustifolium 
3145 bicolor 
589 
590 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Black, G. A. 
47-1496 obovatum 
47-1502 grandiflorum 
47-1889 speciosum 
Black, G. A., Cordeiro, E. & 
Francisco, J. 
52-14649 microearpum 
52-14655 speciosum 
Black, G. A. & Ledoux, P. 
50-10644 sylvestre 
Black, G. A. & Schultes, R. E. 
46-61 grandiflorum 
46-61 (USDA) subincanum 
46-1II grandiflorum 
Boehlmer, F. de 
12229 bicoior 
Bonpland, A. 
s.n. bicoior 
Brenes, A. M. 
12333 bicoior 
British Honduras Forests Con 
servation 
H2192/29 bicoior 
Broadway, W. E. 
8935 angustifolium 
Burchell, W. J. 
9375 grandiflorum 
9467 grandiflorum 
B. W. [Boschwezen] 
1161 velvutinum 
Oalder6n, S. 
627 bicoior 
630 angustifolium 
23610 bicoior 
Camargo, F. C. 
8 microearpum 
2395 speciosum 
Capucho, P. 
397 speciosum 
CardeRosa, B,, Murgueitio, 
P. R. & Bark ley, F. A. 
17C934 bicoior 
Cardona, F, 
2379 subincanum 
Cavalcante, P. 
310 bicoior 
339 obovatum 
937 grandiflorum X obo- 
vatum 
938 subincanum 
Cope, F. W., & Holliday, P. C. 
(see Holliday & Cope) 
Cook, O. F. 
4 bicoior 
Cook, O. F., & Doyle, C. B. 
50 bicoior 
Cook, O. F., & Griggs, R. F. 
756 bicoior 
Cooper, J. J. 
10244 simiarum 
Cooper, G. P., & Slater, G. M. 
242 angustifolium 
Cowan, R. S. 
38164 subincanum 
38186 subincanum 
Cuatrecasas, J. 
6890 subincanum 
7277 subincanum 
7178A subincanum 
14897 cirmolinae 
15336 cirmolinae 
15700 cirmolinae 
16160 bernouillii subsp. capil- 
liferum 
16526A bicoior 
16544 nemorale 
16896 chocoense 
17034 bernouillii subsp. capil- 
liferum 
17034A bernouillii subsp. capil- 
liferum 
17350 bernouillii subsp. capil- 
liferum 
17350A bernouillii subsp. capil- 
liferum 
17503 nemorale 
17738 nemorale 
21291 nemorale 
21339 stipulatum 
Cuatrecasas, J., & Cope, F. W. 
25788 obovatum 
25789 angustifolium 
25790 angustifolium 
25791 mammosum 
25792 simiarum 
25794 simiarum 
25795 bicoior 
25797 obovatum X subincanum 
25800 angustifolium X mammo- 
sum 
25801 grandiflorum 
CUATRECASAS—CACAO AND ITS ALLIES 591 
CUATRECASAS, J., COPE, F. W., & 
Babtlet, B. G. 
25780T grandiflorum 
2578IT grandiflorum 
25782T nemorale 
25783T obovatum 
25784T bicolor 
25785T subincanum 
25786T bicolor 
25787T bicolor 
CuATBECASAS, J., & Le6n, J. 
26515 simiarum 
26515A simiarum 
26516 mammosum 
CUATRECASAS, J., & PAHEDES, A. 
26534 bicolor 
26535 mammosum 
26536 simiarum 
26537 angustifolium 
26538 microcarpum 
ClTATBECASAS, J., & WlLLABD, L, 
26007 nemorale 
26051 nemorale 
26074 chocoense 
26167 gileri 
CuFODONTIS, G. 
92 bicolor j 
599 simiarum 
DaHLGREN, B E., <fc BELLA, E. 
10 microcarpum 
438 grandiflorum 
634 grandiflorum 
733 grandiflorum 
739 grandiflorum 
Davidson, G. W. R. 
s,n. bicolor 
Dawe, M. T. 
83 bicolor 
Dodge, C. W., & Goerger, V. F. 
9420 simiarum 
Dodge, C. W., & Nevebmann 
7164 simiarum 
Dodge, C. W., & Thomas, W. S. 
6399 angustifolium 
Ducke, A. 
100 sylvestre 
103 sylvestre 
265 obovatum 
281 speciosum 
283 microcarpum 
598 grandiflorum 
4878 sylvestre 
6773 microcarpum 
6823 obovatum 
7200 subincanum 
7202 bicolor 
7202B bicolor 
7216 sylvestre 
7397 bicolor 
7638 bicolor 
7679 subincanum 
7704 obovatum 
7822 sylvestre 
7884 speciosum 
7975 speciosum 
10669 sylvestre 
12187 sylvestre 
14734 sylvestre 
14925 subincanum 
16458 grandiflorum 
16464 subincanum 
16466 microcarpum 
21044 obovatum 
21045 microcarpum 
Duque Jabamillo, J. 
1205 bicolor 
4403A bicolor 
Eggebs, H. F. A. 
14244 bicolor 
Ellenbebg, H. 
2551 obovatum 
2565 speciosum 
Escamilla, G. 
s.n. mammosum 
Fagerlind, F.f & Wibom, G. 
2371 bicolor 
FebnAndez P, A. 
2275 subincanum 
Fosbebg, F. R. 
21310 bicolor 
Fbancisco, J. 
1966 grandiflorum 
Fb6eb, B. L, 
20463 sylvestre 
20518 subincanum 
20555 subincanum 
20556 sylvestre 
20625 bicolor 
20645 glaucum 
20646 obovatum 
20648 obovatum X subin- 
canum 
20655 sylvestre 
20750 microcarpum 
20885 glaucum 
20942 glaucum 
592 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
21162 subiocanum 
21482 subincanum 
21556 grandiflorum 
23343 subincanum 
23926 obovatum 
23963 microcarpum 
25554 sylvestre 
26526 speciosum 
28382 sylvestre 
29732 speciosum 
30180 speciosum 
30432 speciosum 
31414 speciosum 
33783 canumanense 
33788 sylvestre 
34663 subincanum 
34949 sylvestre 
Fr6es, R. L., & Filho, J. P. 
29465 sylvestre 
GarcIa Bareiqa, H, 
11178 bicolor 
14203 subincanum 
14224 subincanum 
14253 subincanum 
14380 glaucum 
14416 bicolor 
15139 subincanum 
GarcIa Barrio a, H., Schultes, E. E., 
& Blohm, H. 
16064 subincanum 
Gardner, C. A. 
870 bicolor 
Gentle, P. H, 
3464 bicolor 
Geoffroy, 
s.n. speciosum 
Giler, M. 
162 gileri 
168 gileri 
Giler, M.( & PatiSo, V. M. 
164 gileri 
165 gileri 
166 gileri 
Ginzberger, A. 
802 speciosum 
Ginzberger, A., &. Hagmann, M. 
801 subincanum 
Ginzberoer, A., & Zerner, H. 
800 grandiflorum 
Glaziou, A. F. M. 
9633 speciosum 
9633a subincanum 
9643 grandiflorum 
Goeldi, Andreas 
4225 subincanum 
4226 obovatum 
4228 microcarpum 
Gonggryp, J. W. 
4108 velutinum 
4127 velutinum 
4148 velutinum 
Goudot 
s.n. bicolor 
Guerrero, H. 
26074 chocoense 
Hanburv 
9471 grandiflorum 
Hart, J. H. 
158 bicolor 
5381 augustifolium 
Hoffmannsegg, W. 
s.n. speciosum 
Hold ridge, L. R. 
146 ma mm os um 
5133 gileri 
s.n. bicolor 
Holliday, P. C. 
T 43 subincanum 
T 139 cirmolinae 
T 140 cirmolinae 
T 141 nemorale 
T 142 bernouillii subsp. oapil- 
liferum 
T 143 chocoense 
T 144 chocoense 
T 145 bernouillii subsp. capil- 
Hferum 
T 146 nemorale 
T 147 nemorale 
T 148 nemorale 
T 149 nemorale 
Holliday, P. C., & Bartley, B. G. 
T 163 gileri 
T 165 stipulatum 
T 167 gileri 
T 172 bernouillii subsp. capil- 
liferum 
T 173 nemorale 
T 175 stipulatum 
T 176 chocoense var. bullatum 
T 177 bernouillii subsp. capil- 
liferum 
T 178 bernouillii subsp. capil- 
liferum 
CUATRECASAS—CACAO AND ITS ALLIES 593 
Holliday, P. C., & Cope, F. W. 
T 77 bicolor 
T 79 glaucum 
T 79A glaucum 
T 81 subincanum 
T 90 obovatum 
T 91 subincanum 
T 94 glaucum 
T 95 obovatum 
T 96 glaucum 
T 98 obovatum 
T 101 subincanum 
T 103 subincanum 
T 114 obovatum 
T 115 glaucum 
T 116 subincanum 
T 117 obovaturn X subincanum 
T 118 glaucum 
T 119 obovatum 
T 122 obovatumX subincanum 
T 123 obovatum 
T 124 subincanum 
T 125 microcarpum 
HUBER, H. 
162 subincanum 
1567 speciosum 
1748 speciosum 
4008 grandiflorum 
4295 obovatum 
7081 microcarpum 
Humboldt, A., & Bonpland, A. 
s.n. bicolor 
Idrobo, J. M,, & Schultes, E. E. 
776 subincanum 
INPA (Instituto Nacional db 
Pesquisas da Amazonica, Manaos) 
1966 grandiflorum 
2125 sylvestre 
Jobebt, Dr. 
903 speciosum 
Johnson, H. 
237 bicolor 
Kabsten, G. 
s.n. glaucum 
Killip, E. P., & Smith, A. C. 
30006 bicolor 
30011 grandiflorum 
30320 grandiflorum 
Klug, G. 
87 subincanum 
857 subincanum 
1523 bicolor 
2021 bicolor 
2983 obovatum 
Krukoff, B. 
1080 speciosum 
1117 speciosum 
1274 grandiflorum 
1644 microcarpum 
1668 obovatum 
5295 speciosum 
5383 obovatum 
5759 obovatum 
6203 microcarpum 
6263 obovatum 
6592 microcarpum 
7016 subinpanum 
8226 subincanum 
8275 obovatum 
8280 microcarpum 
9019 bicolor 
Kuhlmann, J. G. 
18110 subincanum 
Labroy 
s.n. grandiflorum 
Lange 
12056 bicolor 
Lehmann, F. C. 
7909 bicolor 
9021 bicolor 
Le6N, J. 
291 mammosum 
937 angustifolium 
1363 mammosum 
3189 si mi arum 
4267 angustifolium 
4832 gileri 
Lindeman, J. C. 
5725 simiarum 
Little, E. L., & Little, R. R. 
9544 subincanum 
9598 bicolor 
Llano, E. 
s.n. bicolor 
L6fez, J. R. 
s.n. angustifolium 
Lucas, A, 
1 bernouillii subsp. asclepi 
adiflorum 
Luetzelbtjbg, Ph. v. ■ 
22007 grandiflorum 
22079 subincanum 
23065 bicolor 
23287 grandiflorum 
23895 bicolor 
594 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Martitts, C. E. P. 
862 bicolor 
863 bicolor 
864 bicolor 
865 bicolor 
871 sylvestre 
872 subincanum 
873 p.p. grandiflorum 
873 p.p. subincanum 
874 grandiflorum 
875 grandiflorum 
876 grandiflorum 
884 microcarpum 
885 microcarpum 
886 microcarpum 
887 sylvestre 
888 sylvestre 
889 sylvestre 
890 sylvestre 
891 sylvestre 
893 subincanum 
894 subincanum 
895 subincanum 
896 subincanum 
897 subincanum 
898 subincanum 
899 subincanum 
900 subincanum 
Observ. 2832 sylvestre 
Observ. 2890 microcarpum 
Matuda, E. 
16690 bicolor 
16733 bicolor 
16840 bicolor 
MexIa, Y. 
7214 bicolor 
Miranda Bastos 
68 subincanum 
Mocxfto, J. M., & Sbss£, M. 
3618 angustifolium 
3620 bicolor 
3621 bicolor 
s.n. angustifolium 
s.n. bicolor 
Monteiro DA Costa 
121 grandiflorum 
Muller, J. V. S. 
s.n. bicolor 
Patin, C. 
s.n. cacao 
PatiRo, V. M. 
22 subincanum 
24 nemorale 
115 chocoense 
116 nemorale 
117 nemorale 
163 grandiflorum 
169 chocoense var. bullatum? 
171 chocoense var. bullatum 
171A chocoense var. bullatum 
171B chocoense var. bullatum 
s.n. speciosum 
Pav6N, J. 
617 bicolor 
201 subincanum 
s.n. bicolor 
s.n, sinuosum 
Gentle, Percy 
3464 bicolor 
Philipson, W. R., Idrobo, J. M., 
& Fernandez, A. 
1552 glaucum 
Pires, J. M. 
136 sylvestre 
4340 bicolor 
4343 grandiflorum X obova- 
tum 
4344 grandiflorum X obova- 
tum 
4345 speciosum X sylvestre 
6575 simiarum 
Pires, J. M., & Black, G. A. 
695 speciosum 
740 speciosum 
742 microcarpum 
743 obovatum 
744 grandiflorum 
746 bicolor 
1414 subincanum 
s.n. obovatum 
Pires, J. M\, Fr6es, R. L., & 
SILVA1 N. T. 
5886 speciosum 
Pires, J. M., Nilo, T., & Silva, A. 
4339 obovatum 
Pittier, H. 
4105 bernouillii 
4194 hylaeum? 
6883 bicolor 
11112 angustifolium 
14016 simiarum 
16142 angustifolium 
s.n. angustifolium 
Pittier, H., & Durand, T. 
3925 simiarum 
8536 angustifolium 
CUATRECASAS—CACAO AND ITS ALLIES 595 
PlTTIER, H., & TONDTJZ, A. 
4074 angustifolium 
Poeppig, E, 
18 bicolor 
1845 obovatum 
2352 p.p. subincanum 
2352 p.p. obovatum 
2746 p.p. obovatum 
2746 p.p. bicolor 
s.n. bicolor 
s.n. obovatum 
Poiteau, A. 
s.n. subincanum 
Preuss, P. 
1381 angustifolium 
Ranoiiel, A. 
195 subincanum 
Reko, B. P. 
6068 bicolor 
Richard, L. C. 
s.n. grandiflorum 
Ribdel, L. 
1373 grandiflorum 
s.n. grandiflorum 
Rivbro 
1836 subincanum 
Romero CastaSeda, R. 
5405 bernouillii subsp. capil- 
liferum 
5500 stipulatum 
s.n. bicolor 
Rufz, H., & Pav6n, J. 
s.n. bicolor 
s.n. sinuosum 
s.n. subincanum 
Rusby, H. H. 
647 speciosum 
654 speciosum 
Sagot, P. 
1206 velutinum 
Sandeman, C. 
2233 grandiflorum 
SCHOMBURQK, R. 
870 p.p. bicolor 
870 p.p. obovatum 
s.n. grandiflorum 
SCHULTES, R. E. 
3471 bicolor 
3922 bicolor 
6536 subincanum 
6921 obovatum 
8065 grandiflorum 
8178 grandiflorum 
8385 subincanum 
12104 subincanum 
Schultes, R, E., Baker, 
R. E. D., & Cabrera, I. 
18552 subincanum 
Schultes, R. E., & Black, G. A. 
8146 grandiflorum 
Schultes, R. E., & Cabrera, I. 
14140 subincanum 
15116 subincanum 
17005 subincanum 
17775 obovatum 
17780 microcarpum 
17781 grandiflorum 
18695 gileri 
Schultes, R. E., & Cordeiro, E. 
6507 speciosum 
Schultes, R. E., & L6pez, F. 
9204 grandiflorum 
Schultes, R. E., 4 Silva, A. 
8066 speciosum 
Schultze-Rhonhof 
2312 glaucum 
Sess£, MociRo, Castillo, & 
Maldonado 
3618 angustifolium 
3620 bicolor 
3621 bicolor 
Siber 
4 grandiflorum 
s.n. speciosum 
Silva, J. F. 
143 subincanum 
155 obovatum 
Silva, A. 
237 subincanum 
317 subincanum 
Siqueiros, R. 
4008 grandiflorum 
Smith, J. Donn, 
6457 8i mi arum 
7313 simiarum 
7731 simiarum 
Snethlage, E. H. 
300 grandiflorum 
10044b speciosum 
Spruce, R, 
97 subincanum 
166 sylvestre 
456 speciosum 
1609 bicolor 
1737 speciosum 
1822 grandiflorum 
CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
s.n. bicolor 
s.n. speciosum 
s.n. sylvestre 
Standley, P. C. 
22317 angustifolium 
36822 simiarum 
37374 bicolor 
37377 simiarum 
79068 angustifolium 
79069 bicolor 
82446 bicolor 
Stern, W., & Chambers, K. 
140 angustifolium 
Steyermark, J. A. 
44941 bicolor 
49317 bicolor 
Stockdale, J. A. 
s.n. bicolor 
StTCHTELEN, N. J. V. 
s.n. simiarum 
Tate, G. H. H. 
944 subincanum 
Tessmann, G. 
3433 obovatum 
4079 bicolor 
4115 subincanum 
4928 sinuosum 
5398 speciosum 
s.n. subincanum 
Tonduz, A. 
4074 angustifolium 
6852 simiarum 
8373 simiarum 
11304 bicolor 
12822 simiarum 
13110 bicolor 
18222 simiarum 
Traill, J. W. R. 
59 subincanum 
60 bicolor 
61 obovatum 
62 glaucum 
s.n. bicolor 
Triana, J. J. 
5333(-3) bicolor 
s.n. bicolor 
Tuekckheim, H. v. 
7824 bicolor 
Ule, E. H. G. 
5030 bicolor 
5637 obovatum 
9609 speciosum 
14448 speciosum 
Weberbaube, A. 
6245 bicolor 
Wedel, H. v. 
681 bernouillii subsp. ascle- 
piadiflorum 
1535 p.p. bernouillii subsp. ascle- 
piadiflorum 
1535 p.p. Lauraceae sp. 
Wickham, H. A. 
s.n. grandiflorum 
Williams, R. 0. 
12121 angustifolium 
Williams, Llewelyn 
161 obovatum 
230 obovatum 
1076 subincanum 
1233 subincanum 
2149 bicolor 
2401 grandiflorum 
3254 subincanum 
3346 bicolor 
5268 obovatum 
9345 bicolor 
15204 subincanum 
15614 grandiflorum 
Collections of Theobroma cacao L. Seen 
Acosta Soils, M., 6332. 10724a. 
Allen, Cyril, 880, 881. 
Asplund, E., 13408, 14464, 14583, 
14788. 
Baker, C. F., 61, 63, 125. 
Baker, R. E. D. & Cope, F. W., 15. 
Banks, s.n. 
Barkley, Araque, & Gomez, 410. 
Bartlett, H. H., 13108. 
Bartley, B. G.f & Holliday, P. C., 51. 
Bernoulli, G., 96, 97, 98. 
Bernoulli, G., & Cario, R., 3150, 3151, 
3152. 
Bertero, C-, 35. 
Blanchet, J. S., 3, 115, 5068, s.n. 
Blanco, 579. 
Bonpland, A., 1102, s.n. 
Box, H. E., 1536. 
Brenes, A. M., 12334. 
Broadway, W. E., 787, 4827, s.n. 
Buchtien, O., 187, s.n. 
Burchell, W. J., 9276. 
Calderdn, S., 107. 
Chevalier, A., s.n. 
Clement, B., 1931. 
Collins, J. H., 15, 16. 
Converse, O., 74. 
Cook, O. F., & Doyle, C. B., 53, 610, 
621, 622, 625, 674, 726. 
Cook, O., & Gilbert, G. B., 1668, 1685. 
Cook, O. F., & Griggs, R. F., 320, 321. 
Cope, F. W,, & Holliday, P., 83, 99, 
102, 104, 105, 107, 111, 127. 
Cuatrecasas et a!., 2555, 7756, 7770, 
13329, 13377, 25802, 25803, 25804, 
25805, 26004, 26005, 26006, 26224, 
26225, 26492, 26493, 26539, 26540, 
26561, 26562, 26563, 26564, 26565. 
Curran, H. M., 122, 163. 
Dahlgren, B. E., et al., 7, 610931. 
Dawe, M. T.( 227. 
Doustan, Dr., s.n. 
Ducke, A., 1095, 12148, 23970, 23976. 
Duque Jaramillo, J., 4411, 4404A. 
Dues, P., 2039, 2900. 
Echevarrfa, 866. 
Emrick, G. M., 14, 
Engel, s.n. 
Espiritu Santo, J., 94. 
Ferreyra, R., 4905. 
Fredholm, A., 3117. 
Frdes, R. L., 20573, 20882, 21484, 
21524, 23925. 
Galeotti, M., 7237. 
Garcia Barriga, H., 8388. 
Garganta, M. de, 717. 
Gentle, P. H., 1740, 3292. 
Glaziou, A. F. M., 9644, 12190. 
Graham, E. H., 500. 
Gregg, 1774. 
Haenke, T., 1533, 2301. 
Hahn, 112. 
Harvey, D., 5215. 
Heller, A. A. & Heller, 726. 
Heyder, H, M., 35. 
Hitchcock, A. E., 449. 
Hinton, G. B., 7531. 
Hodge, W. H., 6715. 
Hohenacker, R. P., 39. 
Holton, I. F., 765. 
Hostmann, W. R., 1, 440. 
Huber, H., 1392, 4392. 
Idrobo, J. M., et al., 784, 940. 
Isert, 87. 
Jack, J. G., 4334. 
Jovert, Dr., 542, 
Jungner, J. R., 79. 
Kappler, A., 1636, s.n. 
Karsten, G., s.n. 
Kellerman, W. A., 4842, 5565, 6045. 
Eidder, N. T., s.n. 
Killip, E. P. <fe Smith, A. C., 29434, 
3022, 33603. 
Klug, G., 926, 2938. 
Krebs, a.n. 
Krukoff, B., 4736, 10661. 
Kuntze, O., s.n. 
Lehmann, F. C., 5641. 
597 
598 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Lemde, A., s.n. 
Le6n, Jorge, 2001. 
Leonard, E. C., 8550, 9249, 9252. 
Levy, P., 1. 
Liebmann, F., 586, 15078, 15079. 
Little, E. L., 6483, 8624. 
Little & Little, 9574. 
Llave, P., s.n. 
Lloyd, 1128. 
Luetzelburg, Ph. v., 2585. 
Lundell, C. L., 2799. 
Macbride, J. P., 5278. 
Martins, C. E. P., 866, 867, 868, 869. 
Matthews, A., 18, s.n., 1653. 
Mell, C. D., 29. 
Mexfa, Y., 6399. 
Miranda, F., 6644, 9299. 
Molina, A., 2346. 
Mulford, 953. 
Myers, J. G., 5829. 
Nadeaud, J., s.n. 
Nelson, E. W., 2490. 
Oca, N. de, 47 bis, 
Orcutt, C. H., 4250. 
Pav6n, J-j 623, s.n. 
P<5rez Arbel&ez, E., 686. 
Philipson, W. R-, et al., 1565, 1569, 
Pierre, 119. 
Pittier, H., 3927, 6615, 11934, 11953, 
s.n. 
Poeppig, E., s.n. 
Poiteau, A., 209, 211, 214, 217. 
Proctor, G. R-, 18348. 
Raunkiaer, Ch., 2525, 2864. 
Reko, B. P., 3393, 4720. 
Richard, L. C., s.n. 
Ricksecker, s.n. 
Rodin, R. J., 594. 
Rose, J. N., 21991. 
Rufz, H. & Pav6n, J., s.n. 
Rusby, H.H., 655. 
Ryan, J., s.n. 
Sagot, P., 52, s.n. 
Sagra, R. de la, 118, 
Salzmann, s.n. 
Sandeman, C., 3382. 
Sargent, F. H., 333. 
Schipp, W. A., 178, 419, s.n. 
Schomburgk, R., s.n. 
Schott, A., s.n. 
Schultes, R. E., et al., 3309, 5858b 
6117, 6667a, 8371, 8524, 8604. 
Scolnick, R., et al., 19An526. 
Sess£ et al., 3619. 
Shafer, J. A., 3428. 
Shannon, W. C., 147. 
Shuttle worth, 1250. 
Sintenis, P., 315, 6370. 
Sneidern, K., Al324. 
Splitzberger, F. L., 1097. 
Standley, P., et al., 19430, 21640 
22699, 25693, 27968, 29673, 30479 
31104, 31384, 44954, 45715, 48644 
52877, 54143, 54879, 55742, 79081, 
82445, 91139. 
Stern, W. 6 al., 171. 
Stevenson, J. A., 116, 3631. 
Steyermark, J. A., 45950, 49218 
54947. 
Swartz, 900, s.n. 
Tessmann, G., 3036. 
Theresa, Prinz., v. Bayern, s.n. 
Thiebout, C., 501. 
Thieme, C., 5156. 
Tonduz, A., 6984, 9928. 
Traill, J. W. R., 58, 63. 
Triana, J. J., 5333, s.n. 
Ule, E. H. G.t 5032. 
Urban, I., 315, 6370. 
Williams, R. S., 806. 
Williams, Llewelyn, 148, 2105, 2349, 
3510, 4160, 5278, 8457, 8981, 9021, 
9022, 9346, 11718, 15869. 
Wilson, P., 162. 
Wright, C., et al., 23X, 2610. 
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CUATRECASAS—CACAO AND ITS ALLIES 603 
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INDEX 
Page references to descriptions or definitions in Boldface. Synonyms in Italics. 
Tribal origins of native names in parentheses 
a-ba-ka-ra (Makuna), 566 
abekarsi (Makuna), 566 
Abroma, 388, 389, 390, 391, 392, 394, 
395, 432, 433, 435, 435 (fig.), 437, 
445 
augusta, 435 (fig.), 443 (fig.), 445, 
588 
augustum, 390 
fastuosum, 390 
nitida, 588 
Acaju, 387 
aligator, 399 
alligator, 399, 400, 402, 513, 516 
Amazonian forastero, 409, 515 
Ambroma, 388 
amelonado, 397, 399, 400, 409, 413, 
501 (fig.), 510, 515, 516, pi. 6 
amelonado amarillo, 394, 402 
amelonado Colorado, 394, 402 
Amygdalae similis Guaiimalensis, 384 
Amygdalis similis guatimalensis, 495 
Amygdalus, 383 
Andropetalum, sect., 421, 422 (fig.), 
425, 431 (map), 451, 452 (key), 
579 
angoleta, 399, 501 (fig.), 508, 516 
a5 (Makuna), 463 
Arriba, 400 
Arvor cacaiifera Americana, 384, 495 
a-so-ya-ee (Piratapuya), 566 
Assonia, 388, 432 
Attalea, 476 
Avellana Mexicana, 384, 495 
Ayenia, 388, 389, 391, 394, 395, 396, 
414, 432, 433 
bacafto, 574, 575 
bacafto dc monte, 547 
bacao, 400, 462, 463, 465, 541, 574, 575 
bacao de monte, 545, 546, 574, 575 
ba-dja-na-hoo (Makuna), 556 
bahla, 400 
balam (Kekchi), 462, 464 
balamati, 462, 464 
balao, 400 
bawk (Maku), 566 
bawk-pom (Maku), 556 
beira de assahyzal, 570 
Uertholletia, 476 
bicco, 488, 489 
bik (Tdrraba), 550 
bizoya, 512 
boldo, 445 
Bombacaceae, 391, 393 
b6o-e (Mirana), 524 
Bouchi-cacao, 485, 486 
Brotobroma aspera, 588 
Bubroma, 388, 390, 396, 400 
(jrandifiorum, 390, 396, 552, 555, 
556 
Gnazuma, 390 
polybotryon, 390 
tomentosum, 390 
Bubroma, sect., 396, 400, 405, 409, 451, 
467, 517, 526 
Bubroma subsect. Glossopetalum, 405, 
526 
Bubroma subsect. Oreantkes, 405, 467 
Bubroma subsect. Telmatocarpus, 405, 
517 
Buettneria, 392, 394 
Buettneriaceae, 442 
Buettnerieae, 392, 394 
Buettneri6es, 437 
Buttneria, 388 
Biittneria, 389, 395, 396 
Biittneriaceae, 391 
Btittnericae, 391, 394, 395, 396 
Buttnerinae, 396 
Byttneria, 388, 432, 433, 435 (fig.), 437 
arguta, 435 (fig.) 
Byttneriaceae, 391, 432, 433, 437 
Byttnerieae, 391, 432, 433, 437 
Byttneri&'s, 433 
Byttnerinae, 433 
cabc?a dc Umbii, 560, 563 
cabega de urubu, 524, 525, 560, 562, 563 
607 
680-695—64 18 
608 INDEX 
cabosse, 428 
cacado dc monte, 512 
cacahoacentli, 383 
cacahoacuahuitl (Nauhatl), 512 
cacahoaquahuitl, 383 
cacahoatl, 379, 383, 512 (Nauhatl) 
cacahuatl (Nauhatl), 379, 512 
cacalto de monte, 574, 575 
Cacao (genus), 383, 384, 385, 386, 387, 
388, 449, 495 
album Peruvianum, 570 
bicolor, 389 
guianensis, 387, 388, 390, 396, 495, 
564, 586, 587, pL 11 
guyanensis, 393 
minor, 385 
minus, 388, 495, 511, 513 
saliva, 387, 388, 401, 495, 510 
sylveslris, 387, 388, 391, 475, 502, 
564, 569, pi. 10 
Theobroma, 495, 511 
Cacao, sect., 393, 406, 409, 495 
Cacao, subsoct., 405, 495 
cacao (common name), 512 
cacao (Kofdn), 466 
cacao amelonado, 512 
cacao azodo, 482, 483 
cacao azul, 474, 475, 470 
cacao biaro, 482 
cacao bianco, 400, 462, 463, 466, 566 
cacao bravo, 463, 466, 525 
cacao calabacillo, 512 
cacao ceniza, 566 
cacao chuncho, 405, 512 
cacao claro, 482 
cacao complex, 406 
cacao eriollo, 394, 404, 512, 513 
cacao d'Anta, 466 
cacao de Castilla, 463 
cacao del pats, 400 
cacao de la India, 533 
cacao de macao, 560, 563 
cacao de mico, 533, 534, 550, 551 
cacao de mono, 550, 551 
cacao de monte, 493, 524, 541, 543, 545, 
546, 566, 569, 574, 577 
cacao de monte bravo, 493 
cacao do matta, 482 
cacao do matto, 482 
cacao dulce, 404, 512 
cacao forastero, 304, 512 
cacao grando de monte, 545 
cacao in(1 to, 541 
j cacao lagarto, 393, 400, 505, 512, 513 
cacao malacayo, 462, 464 
cacao marraco, 463, 466 
cacao meco, 533, 534 
cacao rana, 474, 483, 524, 525, 566 
cacao sacha, 482, 483 
cacao sauvage, 485, 486 
cacao silvestre, 463, 465, 512, 533, 566, 
568, 572, 582 
cacao trinitario, 512 
cacao-hu, 474, 476 
cacaohy, 482, 525 
cacao-i, 482 
cacaolllo, 482 
cacaorana, 483 
cacao-rana, 482, 569 
cacaotlquahuitl (Nauhatl), 512 
cacao-u, 482 
cacao-y, 482 
cacau, 482, 562 
cacau azul, 474, 475, 489 
cacau baju, 463, 466 
cacau bravo, 474, 476, 524, 525 
cacau do Peru, 463, 467 
cacau rana, 474 
cacau silvestre, 516 
cacauatl, 408 
caeauf, 474, 482, 524 
cacau-i, 482, 483 
cacau-rana, 482 
cacau rana, 475, 524 
cacau6, 474, 482 
cacava qualiuitl, 384 
cacavate, 384 
cachu azul, 489 
caco, 512 
cahequa (Tarasc&n), 512 
calabacillo, 394, 397, 398, 399, 400, 402, 
403, 406, 407, 414, 502 (fig.), 510, 
515, 510, 517 
calabacillo amarillo, 394, 397, 401, 402 
calabacillo Colorado, 394, 397, 401, 402 
caocauatzana (Zoque), 512 
Caracas, 399 
carupano, 399 
carupano grando, 399 
carupano legftimo, 399 
carupano mestizo, 399 
carupano parcho, 399 
carupano taparito, 399 
carvu (Kabekara), 462 
cauca, 400 
chocoatl, 379 
INDEX 609 
chocolate, 543 
chocolate de monte, 493, 522, 543, 547, 
572, 574, 575 
chocolatillo, 482, 483 
chucti, 483 
chudechu (Otomi), 512 
coca mono, 533, 534 
cocoa, 512 
coj6n de toro, 400 
Commersonia, 389, 391, 392, 394, 395, 
396, 432, 433, 437 
copuaf, 566 
copu-ai, 560, 563 
copuassti, 556, 557, 558 
Creole, 398 
Crescentia cujete, 463, 556 
criollo, 383, 386, 394, 396, 397, 398, 399, 
400, 401, 402, 403, 405, 406, 407, 
408, 409, 414, 415, 506, 507, 508, 
509, 510, 513, 516, 517 
criollo am&rillo, 396, 400, 401 
criollo caldas, 498 (fig.) 
criollo Caracas, 397 
criollo Colorado, 396, 401 
criollo legltimo, 400 
criollo mestizo, 400 
criollo Venezuela, 507 
cucuh, 512 
cuculat, 512 
cu-Iu-hu (Chokd), 462, 465 
cumacaco, 393, 404 
cumaj6 (Chok6), 547 
cumald, 566, 570 
cundeamor, 394, 397, 399, 516 
cundeamor vars., 402 
cundeamor legltimo, 400 
cundeamor verugoso amarillo, 394, 396 
cundeamor verugoso Colorado, 394, 396 
cundiamor, 498 (fig.), 425 (fig.), 427 
(fig.), 508 
cupai-acti, 556 
cupassd, 556 
cupti do matta, 566 
cupil do matto, 556, 569 
cupd-asstirana, 566 
cupd-assuy, 566 
cupti-curtia, 560, 563 
cupuagti, 466, 556, 557, 558 
cup&-acd, 557 
cupuahy, 566, 570 
cupua-l, 463, 466, 570 
cupuarana, 566, 569 
cupuassA, 463, 466 
cupu-assd, 556, 557, 558 
cupuasU, 556 
cupuf, 566, 570 
cupuhy, 483, 566, 569 
cupurana, 482, 560, 563, 583 
cupu-uassti, 556, 557 
cupuy, 482, 570 
cupuy do igap6, 566, 570 
cupuyh, 482 
curupano grandc, 400 
cushta, 533 
deghy (Otomi), 512 
Dicarpidium, 395 
Diosma, 387 
dol<5 (Doras ke), 512 
Dombeya, 388, 432 
Dombeyaceae, 391 
Dombeyeae, 395 
Dzug-mang-u& (Brunka), 550 
erefa (Guatuso), 462 
Eriolaenae, 391 
Eriolaen6es, 437 
Eutheobroma, Beet., 395, 396, 400, 405, 
409, 458, 495 
Eutheobroma, subsect. Cacao, 405, 495 
Eutheobroma, subsect. Rhytidocarpus, 
405, 458 
farinha, 483 
forastero, 394, 396, 397, 398, 399, 400, 
402, 403, 405, 406, 407, 408, 409, 
414, 506, 515, 516, 517 
forastero amelonado, 401 
forastero amelonado amarillo, 397 
forastero amelonado Colorado, 397 
forastero cundeamor, 401 
forastero ordinary amarillo, 396 
forastero ordinary Colorado, 396 
forastero vars., 402 
Forcipomyia, 432 
Frankliniella parvula, 408, 432 
Glossopetalum, sect., 393,409, 417 (fig.), 
421 (fig.), 422 (fig.), 425, 431 
(map), 438, 451, 452 (key), 453 
(key), 526, 580 
Glossopetalum subsect., 405, 526 
Glossostemon, 389, 391, 392, 394, 432, 
433 
bruguieri, 443, 445 
guandbana, 556 
guayaquil cacao, 399 
Guazuma, 385, 387, 388, 389, 390, 391, 
392, 394, 395, 396, 411, 432, 433, 
435, 435 (fig.), 445, 555 
610 INDEX 
Guazuma—Continued. 
grandiflora, 552 
polybotrya, 443, 445 
tomentosa, 435 (fig.), 588 
ulmirolia, 389, 446, 55a, 5SS 
hd-lia (Tanimuka), 463 
he6-a (Maku), 403 
Helicteraceao, 391 
Hclictereae, 395 
Hcritiera, 442 
Hermannia, 3S8, 395 
Hermannicao, 395 
Her ran ia, 392, 393, 394, 395, 396, 403, 
405, 407, 408, 409, 411, 412, 413, 
432, 433, 435 (koy), 437, 444, 
445 
albiflora, 392, 58S 
balaensis, 588 
camargoana, 444, 588 
cuatrecasana, 425 (fig.), 437 (fie;.), 
444 
gttianensis, 484, 485, 486 
guyanrnsis, 484 
laciniifolia, 392, 588 
mariae, 401, 408, 411, 412, 413, 444, 
588 
nitida, 409, 511, 588 
paraensis, 482 
pulcherrima, 392, 588 
pulcherrima v. pacifica, 437 (fig.), 
443 (fig.), 444 
purpurea, 588 
Hcrrania, s^ct., 395, 396, 400 
Hugonia, 387 
judromaj6 (Choko), 547 
kao-kr«l (Brunka), 512 
kajo (Guatuso), 512 
kako (Mixe), 512 
kau (Tiribf)i 512 
kicob, 512 
kieou, 512 
Klcinhovia, 388, 432 
kno (Penonom6), 512 
ko (Tdrreba), 512 
k6o (Brunka), 512 
kua (Guaimf), 512 
ku-gin (TYirraba), 550 
kuk (Rama), 512 
krilaku (Guatuso), 550 
largarto, 400, 425 (fig.), 500 (fig.), 506, 
514, 516 
largarto amarillo, 514 
largato rojo, 514 
la-na-poc-ta-ma-ca-la-chu-na-ni (Yaku 
na), 463 
Lasiopelaleac. 395 
Tx-ptonyehia, 394, 395, 432, 433, 437 
Licania, 587 
alba, 588 
venosa, 588 
liso amarillo, 394 
liso Colorado, 394 
inacambo, 463, 467 
inachala, 400 
maga (Barasana), 556 
mah-wo-rc (Yukuna), 566 
majambo, 463 
Malvaceae, 393, 395 
Mai vales, 392, 437 
ma-oo-hee-r6e (Kabuyarf), 560, 560 
maraca, 463 
marraco, 463 
Marasmius peruiciosu?, 525, 570, 605 
mate, 463, 556 
ma-wd-roo-da (Kuripaka), 566 
Maxwollia, 394 
maiorca, 428 
mecacahoatl, 383 
Mclhania, 388, 432 
Melochia, 395 
me-tr6-ree-moo-eo (Karihona), 524 
Monilia rorori, 522 
monkey cocoa, 534 
mountain cacao, 464 
nacional Ecuador, 508, 517 
najambu, 463, 467 
Nicaraguan criollo, 402 
no-t6rree-ka (Taniinuka), 566 
nunisup (Rama), 550 
rice-aw (Taniinuka), 556 
Oreanthes, sect., 393, 409, 421 (fig.), 
422 (fig.), 425, 431 (map), 438, 
451, 452 (key), 467 
Oreantkes, subscrt,, 405, 467 
pacxoc, 512 
padamd (Arckuna), 506, 569 
pako kakao, 577 
patachtli, 383 
pataiste, 462, 465 
patas, 463, 466 
patasht, 462, 464 
patashtc, 462, 463, 464 
pa taste, 462, 464 
pataste dc sapo, 462, 464 
pataste simarr6n, 462, 464 
patatlc, 462 
INDEX 611 
pataxte, 462 
pazoli, 464 
pec (Pokonchi), 462 
Pen tape tee, 432 
petaste, 462, 464 
petaxte, 462, 464 
Philippodendrae, 391 
Piptachia, 384 
Polyadelphia Pentandria, 432 
porcelaine, 399 
porcelaine criollo, 516 
porcelaine Java criollo, 514 
poo-hoo (Barasana), 566 
quauhcacahoatl, 383 
ree-ka (Tanimuka), 566 
Rhytidocarpus, sect., 393, 409, 421 
(fig.), 422 (fig.), 425, 438, 451, 
452 (key), 458 
RhytidocarpuSj subsect., 405, 458 
Rulingia, 391, 394, 395, 396, 432, 433, 
437 
sambito, 399, 410 
sangre dc toro, 399 
sapar6n (Estrella), 462 
skar-ub (BribrI), 462 
Sapokaia brasiliensii, 476 
Scaphopetalum, 394, 395, 422, 433, 437 
scar bo (Bribrf), 462 
sor6 (Bribi), 533 
Sterculiaceae, 391, 392, 393, 395, 396, 
432, 433, 437, 442 
Sterculieae, 395, 437, 442 
Surinam, 399 
Telmatocarpus, sect,, 393, 409, 421, 422 
(fig.), 427, 438, 452 (key), 453 
(key), 517, 518 (map) 
Telmatocarpus, subsect., 405, 517 
teta negra, 550 
Thalamiflorae, 392 
Thcobroma, 435 (key), 449, 452 (key), 
455 (key) 
alba, 393, 564, 570, 587, 588 
albijlorum, 588 
album, 395, 398, 405 
angustifolia, 390, 391, 393, 409, 415 
angustifolium, 395, 398, 404, 405, 
407, 414, 427 (fig.), 431 (map), 
440, 441, 443, 446, 454 (key), 456 
(key), 502 (fig.), 526, 528 (fig.), 
531, 533, 539 (fig.), 575, 583, 584, 
585 
angustifolium, X cacao, 583 
angustifolium, X mammosum, 583 
Theobroma—Continued 
asclepiadiflorum, 408, 415, 489, 490, 
491, 492, 493 
aspera, 403, 588 
augusla, 588 
balaensis, 588 
bernouillii, 403, 405, 408, 409, 414, 
431 (map), 440, 443, 453 (key), 
457 (key), 488, 489, 491, 492 
(key), 493 
bernouillii subsp. asclepiadiflorum, 
431 (map), 469 (map), 472 (fig.), 
473 (fig.), 477 (fig.), 492 (key), 
493 
bernouillii subsp. bernouillii, 431 
(map), 469 (map), 472 (fig.), 477 
(fig.), 492 (key) 
bernouillii subsp. capilliferum, 419 
(fig.), 427 (fig.), 429 (fig.), 431 
(map), 469 (map), 472 (fig.), 473 
(fig.), 477 (fig.), 481 (fig.), 492 
(key), 493, pi. 5 
b icolor, 383, 389, 390, 391, 392, 393, 
395, 396, 398, 400, 401, 403, 404, 
405, 407, 409, 411, 412, 413, 414, 
418, 419 (fig.), 425 (fig.), 428, 
437 (fig.), 438, 440, 443, 446, 452, 
456 (key), 458, 459 (fig.), 460 
(map), 479 (fig.), 522, 537 (fig.), 
583, 585, 605 
bicolor X cacao, 411 
bicolor X cacao, 584 
cacao, 417 (fig.), 419 (fig.), 421 
(fig.), 425 (fig.), 427 (fig.), 455 
(key), 481 (fig.), 495, 498 (fig.), 
499 (fig.), 501 (fig.), 503 (fig.), 
512 (key), 508 
cacao sulisp. cacao, 494 (map), 497 
(fig.), 499 (fig.), 512 (key), 513 
cacao subsp. cacao ftna. cacao, 512 
(key) 
cacao subsp. cacao fma. lacando- 
nense, 502 (fig.), 512 (key), 514 
cacao subsp. cacao fma. leiocarpum, 
502 (fig.), 506,512 (key), 514,516 
cacao subsp. cacao fma. pentago- 
Dum, 497 (fig-), 500 (fig.), 512 
(key), 513, 516 
cacao subsp. leiocarpum, 413, 496, 
514, 515 
cacao subsp. pentagona, 496, 513 
cacao subsp. sativa, 496 
612 INDEX 
Theobroma—Continued 
cacao subsp. sphaerocarpum, 494 
(map), 497 (fig.), 501 (fig.), 502 
(fig.), 513 (key), 515, pi. 6 
cacao fma. leiocarpum, 40S, 496, 
515 
cacao fma. pentagonum, 413, 425 
(fig.), 427 (fig.) 
cacao var. leiocarpa, 407, 496, 515 
cacao var. leiocarpitm, 514 
cacao var. typica, 407, 496, 513 
cacao var. typica X v. leiocarpa, 
407, 496 
cacao X inammosum, 584 
cacao X microcarpum, 411 
cacao X obovatum, 411 
cacao X simiarum, 584 
calodesmia, 408, 412, 413, 414, 415, 
486, 488 
Camargoanvm, 413 
camargoanum, 588 
canumanense, 431 (map), 455 (key), 
457 (key), 535 (map), 577, 578, 
579 (fig.) 
capillifera, 414 
capUliferum, 410, 412, 446, 489, 490, 
491, 492, 493 
caribaea, 390, 495, 511 
celtifolia, 389, 588 
chocoense, 412, 421, 421 (fig.), 431 
(map), 454 (key), 458 (key), 529 
(fig.), 535 (map), 538 (fig.), 543, 
545, 546, 549 (fig.), 550, 551, 585 
chocoense var. bullatum, 546 
cirmolinae, 408, 412, 414, 415, 421 
(fig.), 422 (fig.), 423 (fig.), 431 
(map), 446, 453 (key), 458 (key), 
528 (fig.), 531 (fig.), 534, 535 
(map), 536 (fig.), 538 (fig.), 539 
(fig.), pi. 7 
cordata, 460, 467 
ferrugineat 393, 409, 414, 564, 569, 
570, pi. 9 
ferrugineum, 406, 570 
foliis integerrimis, 495 
fo&silium, 587 
gileri, 411, 412, 415, 425 (fig.), 427 
(fig.), 430, 453 (key), 455 (key), 
503 (fig.), 517, 518 (map), 519 
(fig.), 520 (fig.), 521 (fig.), 522 
glauca, 392, 393, 409, 414 
Theobroma—Continued 
glaucum, 391, 395, 398, 405, 419 
(fig.), 431 (map), 443, 443 (fig.), 
444, 451, 453 (key), 457 (key), 
471 (fig.), 473 (fig.), 475 (map), 
477 (fig.), 481 (fig.), 486,488, 493> 
pi. 4 
grandifiora, 409, 415 
grandiflorum, 390, 395, 398, 405, 
407, 408, 411, 412, 413, 414, 421 
(fig.), 425 (fig.), 426 (fig.), 428, 
429 (fig.), 431 (map), 438, 440, 
443, 446, 454, (key), 457 (key), 
474, 526, 533 (fig.), 538 (fig.), 
552, 552 (map), 553 (fig.), 585, 
pi. 8 
grandiflorum X subincanum, 583 
grandiflorum X obovatum, 583 
Guazuma, 386 
guazuma, 588 
guianense, 388, 389, 390, 480, 586, 
587 
guianensis, 390 
hastata, 409, 511, 588 
hylacum, 430, 431, 431 (map), 455 
(key), 458 (key), 502 (fig.), 553 
(fig.), 564 (map), 570, 572, 574, 
575 
integerrima, 389, 495, 511 
Kalagua, 398, 400, 414, 496, 584, 
585, 586 
laciniifolinm, 588 
laeve, 514 
leiocarpa, 393, 410, 446, 495, 514 
leiocarpitm, 395, 404, 405, 406, 407, 
408, 409, 411, 414, 504, 505, 506, 
508, 510, 511, 514, 515 
macrantka, 393, 552, pi. 8 
macrc nthum, 396, 555, 556 
mammosa, 415 
mammosiim, 411, 414, 423, 426, 
(fig.), 431 (map), 452, 456 (key), 
535 (map), 538 (fig.), 567 (fig.), 
573 (fig.), 580, 581, (fig.) 583 
mammosum X simiarum, 583 
mariae, 588 
Mar liana, 391 
Martii, 395, 398, 405, 467, 474 
microcarpum, 391, 393, 395, 398, 
401, 405, 407, 408, 409, 411, 412, 
413, 414, 415, 430, 438, 440, 441, 
443, 444,446,453 (key), 456 (key), 
503 (fig.), 517, 518 (map), 519 
INDEX 613 
Theobroma—Continued 
micro c&rpum—Continued 
(fig.), 521 (fig.), 522, 523, 537 
(fig.) 
montana, 588 
nemorale, 411, 412, 414, 421 (fig.), 
431 (map), 455( key), 457, 473, 
(fig.), 528 (fig-), 529 (fig.), 531, 
564 (map), 567 (figs.), 572, 573 
(fig.), 575 
nemoralis, 415 
nitida, 393, 467, 469, 474, 475 
nitidum, 474, 588 
obovatum, 393, 396, 401, 408, 409, 
411, 412, 413, 414, 415, 431 
(map), 438, 440, 441, 446, 454 
(key), 456 (key), 533 (fig), 537 
(fig.), 552 (map), 553 (fig.), 
559, 561, 562, 583 
obovatum X subincanum, 583 
ovatifolia, 390, 391, 393, 460 
ovotifolium, 390, 396, 398, 400, 463 
Patastle, 463, 464 
pentagona, 393, 402, 403, 410, 414, 
446, 495, 510, 513 
pentagonum, 395, 398, 400, 405, 406, 
409, 410, 411, 414, 504, 505, 506, 
508, 509, 510, 511, 514 
pulckerrimuin, 588 
purpureum, 403, 407, 588 
quinquenervia, 393, 476, pi. 3 
quinquenervium, 396, 480, 483 
sagittata, 409, 410, 511, 588 
saltzmaniana, 393, 410 
Salt zmanniana, 496 
salt zmaPnianum, 504, 511 
salzmavnianum, 395 
sapidutn, 407, 496, 511, 513 
sativa, 401, 410, 496 
sativa var. leucosperma, 410, 496, 
511, 513 
sativa var. melanosperma, 410, 496, 
511 
sativum, 391, 510, 511 
silvestre, 552 
simiarum, 397, 398, 400, 405, 407, 
409, 412, 414, 415, 425 (fig.), 426 
(fig ), 431 (map), 446,454 (key), 
458 (key), 531 (fig.), 538 (fig.), j 
545, 547, 549 (fig.), 550, 561 
(fig.), 583, 584, 585 
sinuata, 575 
Theobroma—Continued 
sinuoxum, 401, 409, 431 (map), 
454 (key), 457 (key), 535 (map), 
573 (fig), 575, 578, 588, pi. 12 
speciosa, 393, 409, 482 
speciosum, 390, 391, 395, 396, 398, 
401, 405, 407, 408, 411, 412, 413, 
414, 425, 427, 431, 425 (fig.), 
427 (fig), 431 (map), 438, 446, 
452 key, 457 key, 467, 468 
(fig.), 471 (fig.), 473 fig., 474, 
475 map, 476, 479 (fig.), 408, 
481 (fig.), 482, 484, 485, 488, 
552, 587, pi. 2, pi. 3. 
speciosum var. coriaceum, 401, 443, 
444, 459 (fig.), 476 
speciosum var. quinquenervia, 476 
speciosum var. Spruceana, 467 
speciosum Xsylvestre, 583 
sphaerocarpa, 401, 403, 410, 496, 
515 
sphaeroearpum, 404, 498 (fig.), 505, 
510 
spruceana, 393, 467, 469 
spruceanum, 396, 405, 408, 412, 
413, 474, 476 
stipulata, 415 
stipulatum, 411, 412, 431 (map), 
453 key, 458 key, 528 (fig), 
537 fig., 546, 550, 551, 531 (fig.), 
535 (map), 539 (fig.), 541, 546, 
585 
subincana, 393 
subincanum, 387, 390, 395, 396, 
398, 401, 405, 407, 408, 412, 
413, 414, 430, 431 (map), 438, 
440, 455 (key), 458 (key), 482, 
521 (fig.), 533 (fig.), 537, 543, 
553 (fig.), 562, 563, 564 (map), 
566, 567 (fig.), 569, 572, 574, 583, 
586, 587, pi. 9, pi. 10, pi. 11 
sylvestre, 390, 395, 398, 401, 405, 
412, 422 (fig.), 431 (map), 437 
(fig.), 438, 440, 441, 452 (key), 
457 (key), 467, 468 (fig.), 469 
(map), 474, 479 (fig.), 559, 562, 
pi. 1 
sylvesiris, 391, 393, 409, 415, 474, 
475,564 
lessmannii, 406, 414, 577, 578, 579, 
pi. 12 
Tessmannii, 575 
614 in] 
Theobroma—Continued 
tomentosa, 396, 588 
undulata, 409, 588 
vclutma, 400 
velutinum, 404, 421 (fig.) t 431 
(map), 452 (key), 456 (key), 459 
(fig.), 471 (fig.), 473 (fig.), 479 
(fig.), 484, 485, 586, 587 
Theobroma, sect., 421 (fig.), 422 (fig.), 
427, 438, 451, 452 (key), 495 
Theobroma sect. Andropetalum, 452 
(key), 579 
Theobroma sect. Cacao, 495 
Theobroma sect. Glossopetalum, 452 
(key), 453 (key), 526 
Theobroma sect. Oreanthes, 452 (key), 
467 
Theobroma sect. Rhytidocarpus, 452 
(key), 458 
Theobroma sect. Telmatocarpus, 452 
(key), 453 (key), 517 
Theobroma sect. Theobroma, 452 (key), 
495 
Theobrominae, 396, 433 
tiger, 463 
Tilia, 388 
tlalcacahoatl, 383 
tlapal, 408 
Tlapalcacauatl, 408 
too-soo (Yauna), 566 
Toxoptera aurantii, 408, 432 
Tribroma, 403, 449 
Tribroma bicolor, 403, 460, 463 
Trinidad criollo, 402 
trinitario, 399, 400, 415, 508, 517 
Trinitario amargo, 400 
Triopteris, 387 
tsiru, (CabGcara), 513 
tsiru, (Bribrl) 512 
Wru-kuru, (Cab6cara) 512 
tutuma, 463 
uchpa-cacao, 566, 570 
uerba (Tdrraba), 462 
Uirub (Bribri), 550 
Uir-ub (Bribrl), 550 
Urubti-acalm, 560, 563 
Venezuelan criollo, 402 
wa-be-ga-ra (Desano), 566 
wa-be-ka-ra (Siriano), 566 
wah-pek-la (Tukano), 566 
wa-k6 (Kubeo), 566 
Waltheria, 395 
wariba, 463 
Wasmannia auropunctata, 408, 432 
wild cacao, 512, 513, 570 
win-cheek (Puinave), 560, 566 
win-cheek-choo-ai (Puinave), 556 
xochicacahoatl, 383 
xocoatl, 379 
yagabizoya (Reko), 512 
yurac-cacao, 566, 570 
V, 5. GOVERNMENT PRINTING OFFICE*