1 Aa 6

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
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.

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

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

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.
ll
409," and Catesb. 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

"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

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.

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,

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

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
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.

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-

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,

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

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

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).

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,

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.

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
very large beans with light-coloured
(6) Var. b. Colorado—Red
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.

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,
(t
solid, dark-colored seeds. T. 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 "
an
1914 d 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:
1
T. bernouillii 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
u
i Originally bo misspelled, although the correct name would have been btrnwUU,**
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.

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

(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-

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
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

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

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-

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
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
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

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
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-
*
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.
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

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.

[Figure 3]
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.

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

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
[Figure 5]
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.
V a
*
i1; ^ J v
i
*
itoiMv
jk /'V I t
I \ 4V
\ \ ^
,r
u H c
■h -.v
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
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.

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

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.
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
vfv
aT,&
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.
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-

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 Byttneriês, 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

[Ficure 8]
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).

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM
436
[Figure 9]
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.

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
1
Contributed bt William L. Stern
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
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, USw Yw FHOw

voucher No.* No. NO.
angustifoUum Cooper Slater Panama Y 10595 3502
242
bernouillii Pitt. Pittier 4105 Panama TJS 30
bicolor H. & B. For. Dept. Br. British F 5632
Hond. Hon-
II. 2192/29(?) duras
tricolor H. & B. "Ford-Brazil Brazil F 22075 6998
397" (?)
bicolor H. & B- LI. Williams Peru F 17804 7008
2149
bicolor H. & B. U. Williams Peru F 18176 7007
3346
cacao Ij. "L. 3225 (via South 5703
Hamburg)" America
cacao L. Vigne 2433 Ghana Kumasi: 6898
K(?)
grandiflorum LI. Williams Peru F 17893 7001
(Willd.) 2401
So hum.
microcarpum Krukoff 6203 Brazil US 36510
Mart.
obovatum LI. Williams Peru F 71232 7010
Ktotzsch ex 161
Bernoulli
obovatum LI. Williams Peru F 17263 7011
Klotzsch ex 230-
Bernoulli
sylvestre Mart. Ducke 103 Brazil Y 21362 7009
subincanum LI. Williams Peru F 17578 7000
Mart.
subincanum LI. Williams Peru F 18144 6999
Mart.
* Abbreviations from W, L. Stern & K\ L* Chambers. The citation of wood specimens and herbarium
vouchers in anatomical research* Taxon* 0: 7-13. I960.
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,

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.

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-

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,

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.

5
Pollen Incompatibility
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.

■
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.
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.
8
Genus Theobroma 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.

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.

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

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

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
• JL * irilArS
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
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
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.

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)
A
Figure 12.—Leaves of Tkeobroma, X H* i 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).
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

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,

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-

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
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,
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
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
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.
Figure 13.—Leaves of Theobroma, X a, speciosutn, from orthotropic and plagiotropic
branches CUS—1031918); b, sylvestre (US—1693105); c, sylvestre (Martius, type).

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
CONTRIBUTIONS FROM THE NATIONAL
470
[Figu&E 14]
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
I /&
.r r• '■ f T ' *
/ if Vi f. •
T■■■ r Jf
It *- ■ | -
i i' ■' ■ r *
G ■ :
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.

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).

"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.

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

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
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).

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,

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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,

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.

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.,
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

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

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.
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 cdtês 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

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
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).

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.

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

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-
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.

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

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
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.

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.
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.).

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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
(t
of seeds each surrounded by their pulp, c, T. cacao subs p. cacao, cultivar criolloM
("Caldas") (Cuatr, 26006 from Colombia), d, Section of same. All X J4*

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."

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,
* .<: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 %•

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
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-

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

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
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

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-
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

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.
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.

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

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):

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.

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,
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

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.

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,

-
k
4
>
«a
H *
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 %.

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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.

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,
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).

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).

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

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
■<■1
i* ■
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

[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),
[Figure 32]
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.

■ '
fif
i , / . ..: y \sr*
?, '.VPr/%
B
&*
t
<Tv. jiv?
"A ■■■■
h "> .. */-.H' : >■ ::-/,■ \U,- ■ ; jC
<y> ;:: y ■/. -,n ..,, V.; r,_^>:'.
i." j : ■ u,.. v ' .■' i
■■'■?■:» V/.■■■■■■ --■ . ■ fr"i-
v ■ ■=- f
■ * .■ « j sr
, K ->: * ■ t
: ■■* \ -:s* '. : , \ . .;
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[Figure 33J
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.
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

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
••••><*
Figure 34.—Thtobroma cirmolinae, flowering and fruiting trunk, X V%.

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

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 :Â, simiarum (Cuatr. 26515A); b, simiarum (Cuatr.
26536); c, ckocoense (Patifio 115); d, cirmolinae (Cuatr. 15336); E, grandiflorum (Cuatr.
25780T); f, mammosum (Cuatr. 26535).

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-
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
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.

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-

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

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

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
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,

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 %•

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<

A
Figure 39.—Leaves of Tkeobroma, X > obovatum (Poeppig 8,n.); b, grandiflorum (Cuatr.
25780T); c, grandiflorum (Killip & Smith 30011); d, kylaeum (Araque & Barkley 18C745);
E, subincanum (Baker 38).

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,

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

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).

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
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).

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

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
;i«W •/.?
I * /■**■■
■H *tPI ■t*'« ,
-.1l1■ ■ - p" 1 Ijfljk»*J|1 ■■1 -U J k"f ■ w1 B-- ■v +
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M»4M H11
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4+f k1I k
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fe:-v"i\ i /- ■■-»/ .iv\ «;J-V ^ ^1"
c 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,
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
563
varzea land, shrub 25 ft. high, "cabeâ 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,
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

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
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).
A
B
* t-
\ % i
% \ \
1.
, ?
\ .
» i
2#W
31
I H" .■ I*f ^ d
j.ki
r-juy
--.\-v ■ ■ :*■. ■ ■.; i ■. ,^ ■*
J v :r
~ ''.
r <v:-;V v|; ; W:: AV.-. rt::* v: -a ^K^VV;!': ^
■ ■ ■ i ■■ i ■ ■ J ■ .
^rT"c!I:îijy■ ■* \\:
1
^ f 1 \\
"*k ' I J ■ Cla AlaJailltl II II| III
Jf" ■ " hi IK" 'W f #«*,"*#+* < ill fl+l i «t 7i;
1 II'mi
: ii*
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
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).

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,
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.
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-
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
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,

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

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.
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

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-

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.

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

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,

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.

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).
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
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
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,

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,

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

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. BAKEB, R. E. D., Baktlet, B. G., &
12392 gileri Holliday, P. C.
12423 gileri 31 subincanum
Allen, P. H. 32a subincanum
3105 grandiflorum BAKEB, R. E. D., & COPE, F, W.
4593 bicolor 2 bicolor
6259 aDgustifolium 3 subincanum
6341 angustifolium 4 subincanum
Appun, C. F. 5 grandiflorum
1 Alba 6 grandiflorum
Araqtje, J., & BAEKLEY, F. A. 7 subincanum
18C745 hylaeum 11 glaucum
Archer, W. A. 11a bicolor
7517 subincanum 12 subincanum
7537 obovatum 13 subincanum
7549 grandiflorum 21 subincanum
7551 microcarpum 25 subincanum
7619 BpecioBum 26 bicolor
7721 speciosum 27 obovatum
7734 grandiflorum 28 microcarpum
7820 subincanum 29 microcarpum
Abplund, E. 30 obovatum
10271 bicolor 31 subincanum
13410 subincanum Bartley, B. G., & Holliday, P. C,
Aublet, J. B. C. F. T 46 grandiflorum
s.n. cacao (fruit) T 47 bicolor
e.n. subincanum T 65 subincanum
BAFOG (Bureau Aob. et Forestier T 66 bicolor
Gutanais) T 68 subincanum
136M velutinum T 69 glaucum
7386 velutinum T 70 glaucum
BAILEY L. H. T 71 subincanum
s.n. angustifolium T 72 subincanum
Baker, C. F. T 74 glaucum
62 grandiflorum T 75 subincanum
421 grandiflorum T 166 hylaeum
2102 angustifolium Bbnoist, R.
Bakeb, R. E. D. 516 velutinum
16 grandiflorum BERNOULLI, G.
33 subincanum 94 bicolor
34 bicolor 95 angustifolium
37 glaucum Bernoulli, G., & Cario, R.
38 subincanum 3188 angustifolium
s.n. angustifolium 3145 bicolor
589
Black, G. A. Cope, F. W., & Holliday, P. C.
47-1496 obovatum (see Holliday & Cope)
47-1502 grandiflorum Cook, O. F.
47-1889 speciosum 4 bicoior
Black, G. A., Cordeiro, E. & Cook, O. F., & Doyle, C. B.
Francisco, J. 50 bicoior
52-14649 microearpum Cook, O. F., & Griggs, R. F.
52-14655 speciosum 756 bicoior
Black, G. A. & Ledoux, P. Cooper, J. J.
50-10644 sylvestre 10244 simiarum
Black, G. A. & Schultes, R. E. Cooper, G. P., & Slater, G. M.
46-61 grandiflorum 242 angustifolium
46-61 (USDA) subincanum Cowan, R. S.
46-1II grandiflorum 38164 subincanum
Boehlmer, F. de 38186 subincanum
12229 bicoior Cuatrecasas, J.
Bonpland, A. 6890 subincanum
s.n. bicoior 7277 subincanum
Brenes, A. M. 7178A subincanum
12333 bicoior 14897 cirmolinae
British Honduras Forests Con 15336 cirmolinae
15700 cirmolinae
servation
16160 bernouillii subsp. capil-

H2192/29 bicoior
liferum
Broadway, W. E.
16526A bicoior
8935 angustifolium
16544 nemorale
Burchell, W. J.
16896 chocoense
9375 grandiflorum
17034 bernouillii subsp. capil-
9467 grandiflorum
liferum
B. W. [Boschwezen] 17034A bernouillii subsp. capil-
1161 velvutinum liferum
Oalder6n, S. 17350 bernouillii subsp. capil-
627 bicoior liferum
630 angustifolium 17350A bernouillii subsp. capil-
23610 bicoior liferum
Camargo, F. C. 17503 nemorale
8 microearpum 17738 nemorale
2395 speciosum 21291 nemorale
Capucho, P. 21339 stipulatum
397 speciosum Cuatrecasas, J., & Cope, F. W.
CardeRosa, B,, Murgueitio, 25788 obovatum
P. R. & Bark ley, F. A. 25789 angustifolium
17C934 bicoior 25790 angustifolium
Cardona, F, 25791 mammosum
2379 subincanum 25792 simiarum
Cavalcante, P. 25794 simiarum
310 bicoior 25795 bicoior
339 obovatum 25797 obovatum X subincanum
937 grandiflorum X obo- 25800 angustifolium X mammo-
vatum sum
938 subincanum 25801 grandiflorum

CUATRECASAS, J., COPE, F. W., & 6823 obovatum
Babtlet, B. G. 7200 subincanum
25780T grandiflorum 7202 bicolor
2578IT grandiflorum 7202B bicolor
25782T nemorale 7216 sylvestre
25783T obovatum 7397 bicolor
25784T bicolor 7638 bicolor
25785T subincanum 7679 subincanum
25786T bicolor 7704 obovatum
25787T bicolor 7822 sylvestre
CuATBECASAS, J., & Le6n, J. 7884 speciosum
26515 simiarum 7975 speciosum
26515A simiarum 10669 sylvestre
26516 mammosum 12187 sylvestre
CUATRECASAS, J., & PAHEDES, A. 14734 sylvestre
26535 mammosum 16458 grandiflorum
26536 simiarum 16464 subincanum
26537 angustifolium 16466 microcarpum
26538 microcarpum 21044 obovatum
ClTATBECASAS, J., & WlLLABD, L, 21045 microcarpum
26007 nemorale Duque Jabamillo, J.
26051 nemorale 1205 bicolor
26074 chocoense 4403A bicolor
26167 gileri Eggebs, H. F. A.
CuFODONTIS, G. 14244 bicolor
92 bicolor j Ellenbebg, H.
599 simiarum 2551 obovatum
DaHLGREN, B E., <fc BELLA, E. 2565 speciosum
10 microcarpum Escamilla, G.
438 grandiflorum s.n. mammosum
634 grandiflorum Fagerlind, F.f & Wibom, G.
733 grandiflorum 2371 bicolor
739 grandiflorum FebnAndez P, A.
Davidson, G. W. R. 2275 subincanum
s,n. bicolor Fosbebg, F. R.
Dawe, M. T. 21310 bicolor
83 bicolor Fbancisco, J.
Dodge, C. W., & Goerger, V. F. 1966 grandiflorum
9420 simiarum Fb6eb, B. L,
Dodge, C. W., & Nevebmann 20463 sylvestre
7164 simiarum 20518 subincanum
Dodge, C. W., & Thomas, W. S. 20555 subincanum
6399 angustifolium 20556 sylvestre
Ducke, A. 20625 bicolor
100 sylvestre 20645 glaucum
103 sylvestre 20646 obovatum
265 obovatum 20648 obovatum X subin-
281 speciosum canum
283 microcarpum 20655 sylvestre
598 grandiflorum 20750 microcarpum
6773 microcarpum 20942 glaucum

21162 subiocanum Goeldi, Andreas
21556 grandiflorum 4226 obovatum
23343 subincanum 4228 microcarpum
23926 obovatum Gonggryp, J. W.
23963 microcarpum 4108 velutinum
25554 sylvestre 4127 velutinum
26526 speciosum 4148 velutinum
28382 sylvestre Goudot
29732 speciosum s.n. bicolor
30180 speciosum Guerrero, H.
30432 speciosum 26074 chocoense
31414 speciosum Hanburv
33783 canumanense 9471 grandiflorum
33788 sylvestre Hart, J. H.
34949 sylvestre 5381 augustifolium
Fr6es, R. L., & Filho, J. P. Hoffmannsegg, W.
29465 sylvestre s.n. speciosum
GarcIa Bareiqa, H, Hold ridge, L. R.
11178 bicolor 146 ma mm os um
14203 subincanum 5133 gileri
14224 subincanum s.n. bicolor
14253 subincanum Holliday, P. C.
14380 glaucum T 43 subincanum
14416 bicolor T 139 cirmolinae
15139 subincanum T 140 cirmolinae
GarcIa Barrio a, H., Schultes, E. E., T 141 nemorale
& Blohm, H. T 142 bernouillii subsp. oapil-
16064 subincanum liferum
Gardner, C. A. T 143 chocoense
870 bicolor T 144 chocoense
Gentle, P. H,
T 145 bernouillii subsp. capil-
3464 bicolor
Hferum
Geoffroy,
T 146 nemorale
s.n. speciosum
T 147 nemorale
Giler, M.
T 148 nemorale
162 gileri
T 149 nemorale
168 gileri
Giler, M.( & PatiSo, V. M. Holliday, P. C., & Bartley, B. G.
164 gileri T 163 gileri
165 gileri T 165 stipulatum
166 gileri T 167 gileri
Ginzberger, A.
802 speciosum
liferum
Ginzberger, A., &. Hagmann, M.
T 173 nemorale
801 subincanum
T 175 stipulatum
Ginzberoer, A., & Zerner, H.
T 176 chocoense var. bullatum
800 grandiflorum

Glaziou, A. F. M.
9633 speciosum liferum
9633a subincanum T 178 bernouillii subsp. capil-
9643 grandiflorum liferum

Holliday, P. C., & Cope, F. W. 2983 obovatum
T 77 bicolor Krukoff, B.
T 79 glaucum 1080 speciosum
T 79A glaucum 1117 speciosum
T 81 subincanum 1274 grandiflorum
T 90 obovatum 1644 microcarpum
T 91 subincanum 1668 obovatum
T 94 glaucum 5295 speciosum
T 95 obovatum 5383 obovatum
T 96 glaucum 5759 obovatum
T 98 obovatum 6203 microcarpum
T 103 subincanum 6592 microcarpum
T 114 obovatum 7016 subinpanum
T 115 glaucum 8226 subincanum
T 117 obovaturn X subincanum 8280 microcarpum
T 118 glaucum 9019 bicolor
T 119 obovatum Kuhlmann, J. G.
T 122 obovatumX subincanum 18110 subincanum
T 123 obovatum Labroy
T 124 subincanum s.n. grandiflorum
T 125 microcarpum Lange
HUBER, H. 12056 bicolor
162 subincanum Lehmann, F. C.
1567 speciosum 7909 bicolor
1748 speciosum 9021 bicolor
4008 grandiflorum Le6N, J.
4295 obovatum 291 mammosum
7081 microcarpum 937 angustifolium
Humboldt, A., & Bonpland, A. 1363 mammosum
s.n. bicolor 3189 si mi arum
Idrobo, J. M,, & Schultes, E. E. 4267 angustifolium
776 subincanum 4832 gileri
INPA (Instituto Nacional db Lindeman, J. C.
Pesquisas da Amazonica, Manaos) 5725 simiarum
1966 grandiflorum Little, E. L., & Little, R. R.
2125 sylvestre 9544 subincanum
Jobebt, Dr. 9598 bicolor
903 speciosum Llano, E.
Johnson, H. s.n. bicolor
237 bicolor L6fez, J. R.
Kabsten, G. s.n. angustifolium
s.n. glaucum
Lucas, A,
Killip, E. P., & Smith, A. C.

1 bernouillii subsp. asclepi
30006 bicolor
adiflorum
30011 grandiflorum
Luetzelbtjbg, Ph. v. ■
30320 grandiflorum
22007 grandiflorum
Klug, G.
22079 subincanum
87 subincanum
23065 bicolor
857 subincanum
1523 bicolor 23287 grandiflorum

Martitts, C. E. P. 115 chocoense
862 bicolor 116 nemorale
863 bicolor 117 nemorale
865 bicolor 169 chocoense var. bullatum?
871 sylvestre 171 chocoense var. bullatum
872 subincanum 171A chocoense var. bullatum
873 p.p. grandiflorum 171B chocoense var. bullatum
873 p.p. subincanum s.n. speciosum
874 grandiflorum Pav6N, J.
876 grandiflorum 201 subincanum
884 microcarpum s.n. bicolor
885 microcarpum s.n, sinuosum
886 microcarpum Gentle, Percy
887 sylvestre 3464 bicolor
888 sylvestre Philipson, W. R., Idrobo, J. M.,
889 sylvestre & Fernandez, A.
891 sylvestre Pires, J. M.
893 subincanum 136 sylvestre
895 subincanum 4343 grandiflorum X obova-
896 subincanum tum
897 subincanum 4344 grandiflorum X obova-
898 subincanum tum
899 subincanum 4345 speciosum X sylvestre
Observ. 2832 sylvestre Pires, J. M., & Black, G. A.
Observ. 2890 microcarpum 695 speciosum
Matuda, E. 740 speciosum
16690 bicolor 742 microcarpum
16733 bicolor 743 obovatum
MexIa, Y. 746 bicolor
Miranda Bastos s.n. obovatum
68 subincanum Pires, J. M\, Fr6es, R. L., &
Mocxfto, J. M., & Sbss£, M. SILVA1 N. T.
3618 angustifolium 5886 speciosum
3620 bicolor Pires, J. M., Nilo, T., & Silva, A.
s.n. angustifolium Pittier, H.
s.n. bicolor 4105 bernouillii
Monteiro DA Costa 4194 hylaeum?
Muller, J. V. S. 11112 angustifolium
s.n. bicolor 14016 simiarum
Patin, C. 16142 angustifolium
s.n. cacao s.n. angustifolium
PatiRo, V. M. Pittier, H., & Durand, T.
24 nemorale 8536 angustifolium

PlTTIER, H., & TONDTJZ, A. 8385 subincanum
Poeppig, E, Schultes, R, E., Baker,
18 bicolor R. E. D., & Cabrera, I.
1845 obovatum 18552 subincanum
2352 p.p. subincanum Schultes, R. E., & Black, G. A.
2352 p.p. obovatum 8146 grandiflorum
2746 p.p. obovatum Schultes, R. E., & Cabrera, I.
2746 p.p. bicolor 14140 subincanum
s.n. bicolor 15116 subincanum
s.n. obovatum 17005 subincanum
Poiteau, A. 17775 obovatum
s.n. subincanum 17780 microcarpum
Preuss, P. 17781 grandiflorum
1381 angustifolium 18695 gileri
Ranoiiel, A. Schultes, R. E., & Cordeiro, E.
195 subincanum 6507 speciosum
Reko, B. P. Schultes, R. E., & L6pez, F.
Richard, L. C. Schultes, R. E., 4 Silva, A.
s.n. grandiflorum 8066 speciosum
Ribdel, L. Schultze-Rhonhof
1373 grandiflorum 2312 glaucum
s.n. grandiflorum Sess£, MociRo, Castillo, &
Rivbro Maldonado
Romero CastaSeda, R. 3620 bicolor
5405 bernouillii subsp. capil- 3621 bicolor
liferum Siber
5500 stipulatum 4 grandiflorum
s.n. bicolor s.n. speciosum
Rufz, H., & Pav6n, J. Silva, J. F.
s.n. bicolor 143 subincanum
s.n. sinuosum 155 obovatum
s.n. subincanum Silva, A.
Rusby, H. H. 237 subincanum
647 speciosum 317 subincanum
654 speciosum Siqueiros, R.
Sagot, P. 4008 grandiflorum
1206 velutinum Smith, J. Donn,
Sandeman, C. 6457 8i mi arum
2233 grandiflorum 7313 simiarum
SCHOMBURQK, R. 7731 simiarum
870 p.p. bicolor Snethlage, E. H.
870 p.p. obovatum 300 grandiflorum
s.n. grandiflorum 10044b speciosum
SCHULTES, R. E. Spruce, R,
3922 bicolor 166 sylvestre
6536 subincanum 456 speciosum
6921 obovatum 1609 bicolor
8065 grandiflorum 1737 speciosum
8178 grandiflorum 1822 grandiflorum

s.n. bicolor 60 bicolor
s.n. speciosum 61 obovatum
s.n. sylvestre 62 glaucum
Standley, P. C. s.n. bicolor
22317 angustifolium Triana, J. J.
36822 simiarum 5333(-3) bicolor
37374 bicolor s.n. bicolor
37377 simiarum Tuekckheim, H. v.
79068 angustifolium 7824 bicolor
79069 bicolor Ule, E. H. G.
Stern, W., & Chambers, K. 5637 obovatum
140 angustifolium 9609 speciosum
Steyermark, J. A. 14448 speciosum
44941 bicolor Weberbaube, A.
Stockdale, J. A. Wedel, H. v.
s.n. bicolor 681 bernouillii subsp. ascle-
StTCHTELEN, N. J. V. piadiflorum
s.n. simiarum 1535 p.p. bernouillii subsp. ascle-
Tate, G. H. H. piadiflorum
944 subincanum 1535 p.p. Lauraceae sp.
Tessmann, G. Wickham, H. A.
3433 obovatum s.n. grandiflorum
4079 bicolor Williams, R. 0.
4928 sinuosum Williams, Llewelyn
5398 speciosum 161 obovatum
s.n. subincanum 230 obovatum
Tonduz, A. 1076 subincanum
6852 simiarum 2149 bicolor
8373 simiarum 2401 grandiflorum
Traill, J. W. R. 15204 subincanum
59 subincanum 15614 grandiflorum

Collections of Theobroma cacao L. Seen
Acosta Soils, M., 6332. 10724a. Dues, P., 2039, 2900.
Allen, Cyril, 880, 881. Echevarrfa, 866.
Asplund, E., 13408, 14464, 14583, Emrick, G. M., 14,
14788. Engel, s.n.
Baker, C. F., 61, 63, 125. Espiritu Santo, J., 94.
Baker, R. E. D. & Cope, F. W., 15. Ferreyra, R., 4905.
Banks, s.n. Fredholm, A., 3117.
Barkley, Araque, & Gomez, 410. Frdes, R. L., 20573, 20882, 21484,
Bartlett, H. H., 13108. 21524, 23925.
Bartley, B. G.f & Holliday, P. C., 51. Galeotti, M., 7237.
Bernoulli, G., 96, 97, 98. Garcia Barriga, H., 8388.
Bernoulli, G., & Cario, R., 3150, 3151, Garganta, M. de, 717.
3152. Gentle, P. H., 1740, 3292.
Bertero, C-, 35. Glaziou, A. F. M., 9644, 12190.
Blanchet, J. S., 3, 115, 5068, s.n. Graham, E. H., 500.
Blanco, 579. Gregg, 1774.
Bonpland, A., 1102, s.n. Haenke, T., 1533, 2301.
Box, H. E., 1536. Hahn, 112.
Brenes, A. M., 12334. Harvey, D., 5215.
Broadway, W. E., 787, 4827, s.n. Heller, A. A. & Heller, 726.
Buchtien, O., 187, s.n. Heyder, H, M., 35.
Burchell, W. J., 9276. Hitchcock, A. E., 449.
Calderdn, S., 107. Hinton, G. B., 7531.
Chevalier, A., s.n. Hodge, W. H., 6715.
Clement, B., 1931. Hohenacker, R. P., 39.
Collins, J. H., 15, 16. Holton, I. F., 765.
Converse, O., 74. Hostmann, W. R., 1, 440.
Cook, O. F., & Doyle, C. B., 53, 610, Huber, H., 1392, 4392.
621, 622, 625, 674, 726. Idrobo, J. M., et al., 784, 940.
Cook, O., & Gilbert, G. B., 1668, 1685. Isert, 87.
Cook, O. F., & Griggs, R. F., 320, 321. Jack, J. G., 4334.
Cope, F. W,, & Holliday, P., 83, 99, Jovert, Dr., 542,
102, 104, 105, 107, 111, 127. Jungner, J. R., 79.
Cuatrecasas et a!., 2555, 7756, 7770, Kappler, A., 1636, s.n.
13329, 13377, 25802, 25803, 25804, Karsten, G., s.n.
25805, 26004, 26005, 26006, 26224, Kellerman, W. A., 4842, 5565, 6045.
26225, 26492, 26493, 26539, 26540, Eidder, N. T., s.n.
26561, 26562, 26563, 26564, 26565. Killip, E. P. <fe Smith, A. C., 29434,
Curran, H. M., 122, 163. 3022, 33603.
Dahlgren, B. E., et al., 7, 610931. Klug, G., 926, 2938.
Dawe, M. T.( 227. Krebs, a.n.
Doustan, Dr., s.n. Krukoff, B., 4736, 10661.
Ducke, A., 1095, 12148, 23970, 23976. Kuntze, O., s.n.
Duque Jaramillo, J., 4411, 4404A. Lehmann, F. C., 5641.
597
Lemde, A., s.n. Sagra, R. de la, 118,
Le6n, Jorge, 2001. Salzmann, s.n.
Leonard, E. C., 8550, 9249, 9252. Sandeman, C., 3382.
Levy, P., 1. Sargent, F. H., 333.
Liebmann, F., 586, 15078, 15079. Schipp, W. A., 178, 419, s.n.
Little, E. L., 6483, 8624. Schomburgk, R., s.n.
Little & Little, 9574. Schott, A., s.n.
Llave, P., s.n. Schultes, R. E., et al., 3309, 5858b
Lloyd, 1128. 6117, 6667a, 8371, 8524, 8604.
Luetzelburg, Ph. v., 2585. Scolnick, R., et al., 19An526.
Lundell, C. L., 2799. Sess£ et al., 3619.
Macbride, J. P., 5278. Shafer, J. A., 3428.
Martins, C. E. P., 866, 867, 868, 869. Shannon, W. C., 147.
Matthews, A., 18, s.n., 1653. Shuttle worth, 1250.
Mell, C. D., 29. Sintenis, P., 315, 6370.
Mexfa, Y., 6399. Sneidern, K., Al324.
Miranda, F., 6644, 9299. Splitzberger, F. L., 1097.
Molina, A., 2346. Standley, P., et al., 19430, 21640
Mulford, 953. 22699, 25693, 27968, 29673, 30479
Myers, J. G., 5829. 31104, 31384, 44954, 45715, 48644
Nadeaud, J., s.n. 52877, 54143, 54879, 55742, 79081,
Nelson, E. W., 2490. 82445, 91139.
Oca, N. de, 47 bis, Stern, W. 6 al., 171.
Orcutt, C. H., 4250. Stevenson, J. A., 116, 3631.
Pav6n, J-j 623, s.n. Steyermark, J. A., 45950, 49218
P<5rez Arbel&ez, E., 686. 54947.
Philipson, W. R-, et al., 1565, 1569, Swartz, 900, s.n.
Pierre, 119. Tessmann, G., 3036.
Pittier, H., 3927, 6615, 11934, 11953, Theresa, Prinz., v. Bayern, s.n.
s.n. Thiebout, C., 501.
Poeppig, E., s.n. Thieme, C., 5156.
Poiteau, A., 209, 211, 214, 217. Tonduz, A., 6984, 9928.
Proctor, G. R-, 18348. Traill, J. W. R., 58, 63.
Raunkiaer, Ch., 2525, 2864. Triana, J. J., 5333, s.n.
Reko, B. P., 3393, 4720.

Ule, E. H. G.t 5032.
Richard, L. C., s.n.

Urban, I., 315, 6370.
Ricksecker, s.n.
Williams, R. S., 806.
Rodin, R. J., 594.
Williams, Llewelyn, 148, 2105, 2349,
Rose, J. N., 21991.
3510, 4160, 5278, 8457, 8981, 9021,
Rufz, H. & Pav6n, J., s.n.
9022, 9346, 11718, 15869.
Rusby, H.H., 655.
Wilson, P., 162.

Ryan, J., s.n.
Sagot, P., 52, s.n. Wright, C., et al., 23X, 2610.

Bibliography
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Addison, G. C., & Miranda Tavareb, T. 1951. Observances sobre as esp£cies
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Allen, Paul H. 1956. The rain forests of Golfo Dulce. Univ. of Florida Press.
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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 balao, 400
abekarsi (Makuna), 566 bawk (Maku), 566
Abroma, 388, 389, 390, 391, 392, 394, bawk-pom (Maku), 556
395, 432, 433, 435, 435 (fig.), 437, beira de assahyzal, 570
445 Uertholletia, 476
augusta, 435 (fig.), 443 (fig.), 445, bicco, 488, 489
588 bik (Tdrraba), 550
augustum, 390 bizoya, 512
fastuosum, 390 boldo, 445
nitida, 588 Bombacaceae, 391, 393
Acaju, 387 b6o-e (Mirana), 524
aligator, 399 Bouchi-cacao, 485, 486
alligator, 399, 400, 402, 513, 516 Brotobroma aspera, 588
Amazonian forastero, 409, 515 Bubroma, 388, 390, 396, 400
Ambroma, 388 (jrandifiorum, 390, 396, 552, 555,
amelonado, 397, 399, 400, 409, 413, 556
501 (fig.), 510, 515, 516, pi. 6 Gnazuma, 390
amelonado amarillo, 394, 402 polybotryon, 390
amelonado Colorado, 394, 402 tomentosum, 390
Amygdalae similis Guaiimalensis, 384 Bubroma, sect., 396, 400, 405, 409, 451,
Amygdalis similis guatimalensis, 495 467, 517, 526
Amygdalus, 383 Bubroma subsect. Glossopetalum, 405,
Andropetalum, sect., 421, 422 (fig.), 526
425, 431 (map), 451, 452 (key), Bubroma subsect. Oreantkes, 405, 467
579 Bubroma subsect. Telmatocarpus, 405,
angoleta, 399, 501 (fig.), 508, 516 517
a5 (Makuna), 463 Buettneria, 392, 394
Arriba, 400 Buettneriaceae, 442
Arvor cacaiifera Americana, 384, 495 Buettnerieae, 392, 394
a-so-ya-ee (Piratapuya), 566 Buettneri6es, 437
Assonia, 388, 432 Buttneria, 388
Attalea, 476 Biittneria, 389, 395, 396
Avellana Mexicana, 384, 495 Biittneriaceae, 391
Ayenia, 388, 389, 391, 394, 395, 396, Btittnericae, 391, 394, 395, 396
414, 432, 433 Buttnerinae, 396
bacafto, 574, 575 Byttneria, 388, 432, 433, 435 (fig.), 437
bacafto dc monte, 547 arguta, 435 (fig.)
bacao, 400, 462, 463, 465, 541, 574, 575 Byttneriaceae, 391, 432, 433, 437
bacao de monte, 545, 546, 574, 575 Byttnerieae, 391, 432, 433, 437
ba-dja-na-hoo (Makuna), 556 Byttneri&'s, 433
bahla, 400 Byttnerinae, 433
balam (Kekchi), 462, 464 cabc?a dc Umbii, 560, 563
balamati, 462, 464 cabega de urubu, 524, 525, 560, 562, 563
607
680-695—64 18
608 INDEX
cabosse, 428 j cacao lagarto, 393, 400, 505, 512, 513
cacado dc monte, 512 cacao malacayo, 462, 464
cacahoacentli, 383 cacao marraco, 463, 466
cacahoacuahuitl (Nauhatl), 512 cacao meco, 533, 534
cacahoaquahuitl, 383 cacao rana, 474, 483, 524, 525, 566
cacahoatl, 379, 383, 512 (Nauhatl) cacao sacha, 482, 483
cacahuatl (Nauhatl), 379, 512 cacao sauvage, 485, 486
cacalto de monte, 574, 575 cacao silvestre, 463, 465, 512, 533, 566,
Cacao (genus), 383, 384, 385, 386, 387, 568, 572, 582
388, 449, 495 cacao trinitario, 512
album Peruvianum, 570 cacao-hu, 474, 476
bicolor, 389 cacaohy, 482, 525
guianensis, 387, 388, 390, 396, 495, cacao-i, 482
564, 586, 587, pL 11 cacaolllo, 482
guyanensis, 393 cacaorana, 483
minor, 385 cacao-rana, 482, 569
minus, 388, 495, 511, 513 cacaotlquahuitl (Nauhatl), 512
saliva, 387, 388, 401, 495, 510 cacao-u, 482
sylveslris, 387, 388, 391, 475, 502, cacao-y, 482
564, 569, pi. 10 cacau, 482, 562
Theobroma, 495, 511 cacau azul, 474, 475, 489
Cacao, sect., 393, 406, 409, 495 cacau baju, 463, 466
Cacao, subsoct., 405, 495 cacau bravo, 474, 476, 524, 525
cacao (common name), 512 cacau do Peru, 463, 467
cacao (Kofdn), 466 cacau rana, 474
cacao amelonado, 512 cacau silvestre, 516
cacao azodo, 482, 483 cacauatl, 408
cacao azul, 474, 475, 470 caeauf, 474, 482, 524
cacao biaro, 482 cacau-i, 482, 483
cacao bianco, 400, 462, 463, 466, 566 cacau-rana, 482
cacao bravo, 463, 466, 525 cacau rana, 475, 524
cacao calabacillo, 512 cacau6, 474, 482
cacao ceniza, 566 cacava qualiuitl, 384
cacao chuncho, 405, 512 cacavate, 384
cacao claro, 482 cachu azul, 489
cacao complex, 406 caco, 512
cacao eriollo, 394, 404, 512, 513 cahequa (Tarasc&n), 512
cacao d'Anta, 466 calabacillo, 394, 397, 398, 399, 400, 402,
cacao de Castilla, 463 403, 406, 407, 414, 502 (fig.), 510,
cacao del pats, 400 515, 510, 517
cacao de la India, 533 calabacillo amarillo, 394, 397, 401, 402
cacao de macao, 560, 563 calabacillo Colorado, 394, 397, 401, 402
cacao de mico, 533, 534, 550, 551 caocauatzana (Zoque), 512
cacao de mono, 550, 551 Caracas, 399
cacao de monte, 493, 524, 541, 543, 545, carupano, 399
546, 566, 569, 574, 577 carupano grando, 399
cacao de monte bravo, 493 carupano legftimo, 399
cacao do matta, 482 carupano mestizo, 399
cacao do matto, 482 carupano parcho, 399
cacao dulce, 404, 512 carupano taparito, 399
cacao forastero, 304, 512 carvu (Kabekara), 462
cacao grando de monte, 545 cauca, 400
cacao in(1 to, 541 chocoatl, 379

INDEX 609
chocolate, 543 cupu-assd, 556, 557, 558
chocolate de monte, 493, 522, 543, 547, cupuasU, 556
572, 574, 575 cupuf, 566, 570
chocolatillo, 482, 483 cupuhy, 483, 566, 569
chucti, 483 cupurana, 482, 560, 563, 583
chudechu (Otomi), 512 cupu-uassti, 556, 557
coca mono, 533, 534 cupuy, 482, 570
cocoa, 512 cupuy do igap6, 566, 570
coj6n de toro, 400 cupuyh, 482
Commersonia, 389, 391, 392, 394, 395, curupano grandc, 400
396, 432, 433, 437 cushta, 533
copuaf, 566 deghy (Otomi), 512
copu-ai, 560, 563 Dicarpidium, 395
copuassti, 556, 557, 558 Diosma, 387
Creole, 398 dol<5 (Doras ke), 512
Crescentia cujete, 463, 556 Dombeya, 388, 432
criollo, 383, 386, 394, 396, 397, 398, 399, Dombeyaceae, 391
400, 401, 402, 403, 405, 406, 407, Dombeyeae, 395
408, 409, 414, 415, 506, 507, 508, Dzug-mang-u& (Brunka), 550
509, 510, 513, 516, 517 erefa (Guatuso), 462
criollo am&rillo, 396, 400, 401 Eriolaenae, 391
criollo caldas, 498 (fig.) Eriolaen6es, 437
criollo Caracas, 397 Eutheobroma, Beet., 395, 396, 400, 405,
criollo Colorado, 396, 401 409, 458, 495
criollo legltimo, 400 Eutheobroma, subsect. Cacao, 405, 495
criollo mestizo, 400 Eutheobroma, subsect. Rhytidocarpus,
criollo Venezuela, 507 405, 458
cucuh, 512 farinha, 483
cuculat, 512 forastero, 394, 396, 397, 398, 399, 400,
cu-Iu-hu (Chokd), 462, 465 402, 403, 405, 406, 407, 408, 409,
cumacaco, 393, 404 414, 506, 515, 516, 517
cumaj6 (Chok6), 547 forastero amelonado, 401
cumald, 566, 570 forastero amelonado amarillo, 397
cundeamor, 394, 397, 399, 516 forastero amelonado Colorado, 397
cundeamor vars., 402 forastero cundeamor, 401
cundeamor legltimo, 400 forastero ordinary amarillo, 396
cundeamor verugoso amarillo, 394, 396 forastero ordinary Colorado, 396
cundeamor verugoso Colorado, 394, 396 forastero vars., 402
cundiamor, 498 (fig.), 425 (fig.), 427 Forcipomyia, 432
(fig.), 508 Frankliniella parvula, 408, 432
cupai-acti, 556 Glossopetalum, sect., 393,409, 417 (fig.),
cupassd, 556 421 (fig.), 422 (fig.), 425, 431
cupti do matta, 566 (map), 438, 451, 452 (key), 453
cupil do matto, 556, 569 (key), 526, 580
cupd-asstirana, 566 Glossopetalum subsect., 405, 526
cupd-assuy, 566 Glossostemon, 389, 391, 392, 394, 432,
cupti-curtia, 560, 563 433
cupuagti, 466, 556, 557, 558 bruguieri, 443, 445
cup&-acd, 557 guandbana, 556
cupuahy, 566, 570 guayaquil cacao, 399
cupua-l, 463, 466, 570 Guazuma, 385, 387, 388, 389, 390, 391,
cupuarana, 566, 569 392, 394, 395, 396, 411, 432, 433,
cupuassA, 463, 466 435, 435 (fig.), 445, 555

610 INDEX
Guazuma—Continued. la-na-poc-ta-ma-ca-la-chu-na-ni (Yaku
grandiflora, 552 na), 463
polybotrya, 443, 445 Lasiopelaleac. 395
tomentosa, 435 (fig.), 588 Tx-ptonyehia, 394, 395, 432, 433, 437
ulmirolia, 389, 446, 55a, 5SS Licania, 587
hd-lia (Tanimuka), 463 alba, 588
he6-a (Maku), 403 venosa, 588
Helicteraceao, 391 liso amarillo, 394
Hclictereae, 395 liso Colorado, 394
Hcritiera, 442 inacambo, 463, 467
Hermannia, 3S8, 395 inachala, 400
Hermannicao, 395 maga (Barasana), 556
Her ran ia, 392, 393, 394, 395, 396, 403, mah-wo-rc (Yukuna), 566
405, 407, 408, 409, 411, 412, 413, majambo, 463
432, 433, 435 (koy), 437, 444, Malvaceae, 393, 395
445 Mai vales, 392, 437
albiflora, 392, 58S ma-oo-hee-r6e (Kabuyarf), 560, 560
balaensis, 588 maraca, 463
camargoana, 444, 588 marraco, 463
cuatrecasana, 425 (fig.), 437 (fie;.), Marasmius peruiciosu?, 525, 570, 605
444 mate, 463, 556
gttianensis, 484, 485, 486 ma-wd-roo-da (Kuripaka), 566
guyanrnsis, 484 Maxwollia, 394
laciniifolia, 392, 588 maiorca, 428
mariae, 401, 408, 411, 412, 413, 444, mecacahoatl, 383
588 Mclhania, 388, 432
nitida, 409, 511, 588 Melochia, 395
paraensis, 482 me-tr6-ree-moo-eo (Karihona), 524
pulcherrima, 392, 588 Monilia rorori, 522
pulcherrima v. pacifica, 437 (fig.), monkey cocoa, 534
443 (fig.), 444 mountain cacao, 464
purpurea, 588 nacional Ecuador, 508, 517
Hcrrania, s^ct., 395, 396, 400 najambu, 463, 467
Hugonia, 387 Nicaraguan criollo, 402
judromaj6 (Choko), 547 no-t6rree-ka (Taniinuka), 566
kao-kr«l (Brunka), 512 nunisup (Rama), 550
kajo (Guatuso), 512 rice-aw (Taniinuka), 556
kako (Mixe), 512 Oreanthes, sect., 393, 409, 421 (fig.),
kau (Tiribf)i 512 422 (fig.), 425, 431 (map), 438,
kicob, 512 451, 452 (key), 467
kieou, 512 Oreantkes, subscrt,, 405, 467
Klcinhovia, 388, 432 pacxoc, 512
kno (Penonom6), 512 padamd (Arckuna), 506, 569
ko (Tdrreba), 512 pako kakao, 577
k6o (Brunka), 512 patachtli, 383
kua (Guaimf), 512 pataiste, 462, 465
ku-gin (TYirraba), 550 patas, 463, 466
kuk (Rama), 512 patasht, 462, 464
krilaku (Guatuso), 550 patashtc, 462, 463, 464
largarto, 400, 425 (fig.), 500 (fig.), 506, pa taste, 462, 464
514, 516 pataste dc sapo, 462, 464
largarto amarillo, 514 pataste simarr6n, 462, 464
largato rojo, 514 patatlc, 462

INDEX 611
pataxte, 462 Theobroma—Continued
pazoli, 464 asclepiadiflorum, 408, 415, 489, 490,
pec (Pokonchi), 462 491, 492, 493
Pen tape tee, 432 aspera, 403, 588
petaste, 462, 464 augusla, 588
petaxte, 462, 464 balaensis, 588
Philippodendrae, 391 bernouillii, 403, 405, 408, 409, 414,
Piptachia, 384 431 (map), 440, 443, 453 (key),
Polyadelphia Pentandria, 432 457 (key), 488, 489, 491, 492
porcelaine, 399 (key), 493
porcelaine criollo, 516 bernouillii subsp. asclepiadiflorum,
porcelaine Java criollo, 514 431 (map), 469 (map), 472 (fig.),
poo-hoo (Barasana), 566 473 (fig.), 477 (fig.), 492 (key),
quauhcacahoatl, 383 493
ree-ka (Tanimuka), 566 bernouillii subsp. bernouillii, 431
Rhytidocarpus, sect., 393, 409, 421 (map), 469 (map), 472 (fig.), 477
(fig.), 422 (fig.), 425, 438, 451, (fig.), 492 (key)
452 (key), 458 bernouillii subsp. capilliferum, 419
RhytidocarpuSj subsect., 405, 458 (fig.), 427 (fig.), 429 (fig.), 431
Rulingia, 391, 394, 395, 396, 432, 433, (map), 469 (map), 472 (fig.), 473
437 (fig.), 477 (fig.), 481 (fig.), 492
sambito, 399, 410 (key), 493, pi. 5
sangre dc toro, 399 b icolor, 383, 389, 390, 391, 392, 393,
sapar6n (Estrella), 462 395, 396, 398, 400, 401, 403, 404,
skar-ub (BribrI), 462 405, 407, 409, 411, 412, 413, 414,
Sapokaia brasiliensii, 476 418, 419 (fig.), 425 (fig.), 428,
Scaphopetalum, 394, 395, 422, 433, 437 437 (fig.), 438, 440, 443, 446, 452,
scar bo (Bribrf), 462 456 (key), 458, 459 (fig.), 460
sor6 (Bribi), 533 (map), 479 (fig.), 522, 537 (fig.),
Sterculiaceae, 391, 392, 393, 395, 396, 583, 585, 605
432, 433, 437, 442 bicolor X cacao, 411
Sterculieae, 395, 437, 442

bicolor X cacao, 584
Surinam, 399
cacao, 417 (fig.), 419 (fig.), 421
Telmatocarpus, sect,, 393, 409, 421, 422
(fig.), 425 (fig.), 427 (fig.), 455
(fig.), 427, 438, 452 (key), 453

(key), 481 (fig.), 495, 498 (fig.),
(key), 517, 518 (map)
499 (fig.), 501 (fig.), 503 (fig.),
Telmatocarpus, subsect., 405, 517
512 (key), 508
teta negra, 550
cacao sulisp. cacao, 494 (map), 497
Thalamiflorae, 392
(fig.), 499 (fig.), 512 (key), 513
Thcobroma, 435 (key), 449, 452 (key),
cacao subsp. cacao ftna. cacao, 512
455 (key)
(key)
alba, 393, 564, 570, 587, 588
cacao subsp. cacao fma. lacando-
albijlorum, 588
nense, 502 (fig.), 512 (key), 514
album, 395, 398, 405
cacao subsp. cacao fma. leiocarpum,

angustifolia, 390, 391, 393, 409, 415
502 (fig.), 506,512 (key), 514,516

angustifolium, 395, 398, 404, 405,
407, 414, 427 (fig.), 431 (map), cacao subsp. cacao fma. pentago-
Dum, 497 (fig-), 500 (fig.), 512

440, 441, 443, 446, 454 (key), 456
(key), 502 (fig.), 526, 528 (fig.), (key), 513, 516
531, 533, 539 (fig.), 575, 583, 584, cacao subsp. leiocarpum, 413, 496,
585 514, 515
angustifolium, X cacao, 583 cacao subsp. pentagona, 496, 513
angustifolium, X mammosum, 583 cacao subsp. sativa, 496

612 INDEX
Theobroma—Continued Theobroma—Continued
cacao subsp. sphaerocarpum, 494 glaucum, 391, 395, 398, 405, 419
(map), 497 (fig.), 501 (fig.), 502 (fig.), 431 (map), 443, 443 (fig.),
(fig.), 513 (key), 515, pi. 6 444, 451, 453 (key), 457 (key),
cacao fma. leiocarpum, 40S, 496, 471 (fig.), 473 (fig.), 475 (map),
515 477 (fig.), 481 (fig.), 486,488, 493>
cacao fma. pentagonum, 413, 425 pi. 4
(fig.), 427 (fig.) grandifiora, 409, 415
cacao var. leiocarpa, 407, 496, 515 grandiflorum, 390, 395, 398, 405,
cacao var. leiocarpitm, 514 407, 408, 411, 412, 413, 414, 421
cacao var. typica, 407, 496, 513 (fig.), 425 (fig.), 426 (fig.), 428,
cacao var. typica X v. leiocarpa, 429 (fig.), 431 (map), 438, 440,
407, 496 443, 446, 454, (key), 457 (key),
cacao X inammosum, 584 474, 526, 533 (fig.), 538 (fig.),
cacao X microcarpum, 411 552, 552 (map), 553 (fig.), 585,
cacao X obovatum, 411 pi. 8
cacao X simiarum, 584 grandiflorum X subincanum, 583
calodesmia, 408, 412, 413, 414, 415, grandiflorum X obovatum, 583
486, 488 Guazuma, 386
Camargoanvm, 413 guazuma, 588
camargoanum, 588 guianense, 388, 389, 390, 480, 586,
canumanense, 431 (map), 455 (key), 587
457 (key), 535 (map), 577, 578, guianensis, 390
579 (fig.) hastata, 409, 511, 588
capillifera, 414 hylacum, 430, 431, 431 (map), 455
capUliferum, 410, 412, 446, 489, 490, (key), 458 (key), 502 (fig.), 553
491, 492, 493 (fig.), 564 (map), 570, 572, 574,
caribaea, 390, 495, 511 575
celtifolia, 389, 588 integerrima, 389, 495, 511
chocoense, 412, 421, 421 (fig.), 431 Kalagua, 398, 400, 414, 496, 584,
(map), 454 (key), 458 (key), 529 585, 586
(fig.), 535 (map), 538 (fig.), 543, laciniifolinm, 588
laeve, 514
545, 546, 549 (fig.), 550, 551, 585
leiocarpa, 393, 410, 446, 495, 514

chocoense var. bullatum, 546
leiocarpitm, 395, 404, 405, 406, 407,
cirmolinae, 408, 412, 414, 415, 421
408, 409, 411, 414, 504, 505, 506,
(fig.), 422 (fig.), 423 (fig.), 431
508, 510, 511, 514, 515
(map), 446, 453 (key), 458 (key),
macrantka, 393, 552, pi. 8
528 (fig.), 531 (fig.), 534, 535
macrc nthum, 396, 555, 556
(map), 536 (fig.), 538 (fig.), 539

mammosa, 415
(fig.), pi. 7 mammosiim, 411, 414, 423, 426,
cordata, 460, 467 (fig.), 431 (map), 452, 456 (key),
ferrugineat 393, 409, 414, 564, 569, 535 (map), 538 (fig.), 567 (fig.),
570, pi. 9 573 (fig.), 580, 581, (fig.) 583
mammosum X simiarum, 583

ferrugineum, 406, 570
mariae, 588
foliis integerrimis, 495
Mar liana, 391
fo&silium, 587
Martii, 395, 398, 405, 467, 474
gileri, 411, 412, 415, 425 (fig.), 427
microcarpum, 391, 393, 395, 398,
(fig.), 430, 453 (key), 455 (key),
401, 405, 407, 408, 409, 411, 412,
503 (fig.), 517, 518 (map), 519

413, 414, 415, 430, 438, 440, 441,
(fig.), 520 (fig.), 521 (fig.), 522 443, 444,446,453 (key), 456 (key),
glauca, 392, 393, 409, 414 503 (fig.), 517, 518 (map), 519
INDEX 613
Theobroma—Continued Theobroma—Continued
micro c&rpum—Continued sinuoxum, 401, 409, 431 (map),
(fig.), 521 (fig.), 522, 523, 537 454 (key), 457 (key), 535 (map),
(fig.) 573 (fig), 575, 578, 588, pi. 12
montana, 588 speciosa, 393, 409, 482
nemorale, 411, 412, 414, 421 (fig.), speciosum, 390, 391, 395, 396, 398,
431 (map), 455( key), 457, 473, 401, 405, 407, 408, 411, 412, 413,
(fig.), 528 (fig-), 529 (fig.), 531, 414, 425, 427, 431, 425 (fig.),
564 (map), 567 (figs.), 572, 573 427 (fig), 431 (map), 438, 446,
(fig.), 575 452 key, 457 key, 467, 468
nemoralis, 415 (fig.), 471 (fig.), 473 fig., 474,
nitida, 393, 467, 469, 474, 475 475 map, 476, 479 (fig.), 408,
nitidum, 474, 588 481 (fig.), 482, 484, 485, 488,
obovatum, 393, 396, 401, 408, 409, 552, 587, pi. 2, pi. 3.
411, 412, 413, 414, 415, 431 speciosum var. coriaceum, 401, 443,
(map), 438, 440, 441, 446, 454 444, 459 (fig.), 476
(key), 456 (key), 533 (fig), 537 speciosum var. quinquenervia, 476
(fig.), 552 (map), 553 (fig.), speciosum var. Spruceana, 467
559, 561, 562, 583 speciosum Xsylvestre, 583
obovatum X subincanum, 583 sphaerocarpa, 401, 403, 410, 496,
ovatifolia, 390, 391, 393, 460 515
ovotifolium, 390, 396, 398, 400, 463 sphaeroearpum, 404, 498 (fig.), 505,
Patastle, 463, 464 510
pentagona, 393, 402, 403, 410, 414, spruceana, 393, 467, 469
446, 495, 510, 513

spruceanum, 396, 405, 408, 412,
pentagonum, 395, 398, 400, 405, 406,

413, 474, 476
409, 410, 411, 414, 504, 505, 506,

stipulata, 415
508, 509, 510, 511, 514

stipulatum, 411, 412, 431 (map),
pulckerrimuin, 588
453 key, 458 key, 528 (fig),
purpureum, 403, 407, 588
537 fig., 546, 550, 551, 531 (fig.),
quinquenervia, 393, 476, pi. 3
535 (map), 539 (fig.), 541, 546,
quinquenervium, 396, 480, 483
585
sagittata, 409, 410, 511, 588
subincana, 393
saltzmaniana, 393, 410
subincanum, 387, 390, 395, 396,
Salt zmanniana, 496
398, 401, 405, 407, 408, 412,
salt zmaPnianum, 504, 511
413, 414, 430, 431 (map), 438,
salzmavnianum, 395
440, 455 (key), 458 (key), 482,
sapidutn, 407, 496, 511, 513 521 (fig.), 533 (fig.), 537, 543,
sativa, 401, 410, 496 553 (fig.), 562, 563, 564 (map),
sativa var. leucosperma, 410, 496, 566, 567 (fig.), 569, 572, 574, 583,
511, 513 586, 587, pi. 9, pi. 10, pi. 11
sativa var. melanosperma, 410, 496, sylvestre, 390, 395, 398, 401, 405,
511 412, 422 (fig.), 431 (map), 437
sativum, 391, 510, 511 (fig.), 438, 440, 441, 452 (key),
silvestre, 552 457 (key), 467, 468 (fig.), 469
(map), 474, 479 (fig.), 559, 562,

simiarum, 397, 398, 400, 405, 407,
pi. 1
409, 412, 414, 415, 425 (fig.), 426
(fig ), 431 (map), 446,454 (key), sylvesiris, 391, 393, 409, 415, 474,
475,564
458 (key), 531 (fig.), 538 (fig.), j
545, 547, 549 (fig.), 550, 561 lessmannii, 406, 414, 577, 578, 579,
(fig.), 583, 584, 585 pi. 12
sinuata, 575 Tessmannii, 575

614 in]
Theobroma—Continued Tribroma, 403, 449
tomentosa, 396, 588 Tribroma bicolor, 403, 460, 463
undulata, 409, 588 Trinidad criollo, 402
vclutma, 400 trinitario, 399, 400, 415, 508, 517
velutinum, 404, 421 (fig.) t 431 Trinitario amargo, 400
(map), 452 (key), 456 (key), 459 Triopteris, 387
(fig.), 471 (fig.), 473 (fig.), 479 tsiru, (CabGcara), 513
(fig.), 484, 485, 586, 587 tsiru, (Bribrl) 512
Theobroma, sect., 421 (fig.), 422 (fig.), Wru-kuru, (Cab6cara) 512
427, 438, 451, 452 (key), 495 tutuma, 463
Theobroma sect. Andropetalum, 452 uchpa-cacao, 566, 570
(key), 579 uerba (Tdrraba), 462
Theobroma sect. Cacao, 495 Uirub (Bribri), 550
Theobroma sect. Glossopetalum, 452 Uir-ub (Bribrl), 550
(key), 453 (key), 526 Urubti-acalm, 560, 563
Theobroma sect. Oreanthes, 452 (key), Venezuelan criollo, 402
467 wa-be-ga-ra (Desano), 566
Theobroma sect. Rhytidocarpus, 452 wa-be-ka-ra (Siriano), 566
(key), 458 wah-pek-la (Tukano), 566
Theobroma sect. Telmatocarpus, 452 wa-k6 (Kubeo), 566
(key), 453 (key), 517 Waltheria, 395
Theobroma sect. Theobroma, 452 (key),

wariba, 463
495
Wasmannia auropunctata, 408, 432
Theobrominae, 396, 433
wild cacao, 512, 513, 570
tiger, 463
win-cheek (Puinave), 560, 566
Tilia, 388
win-cheek-choo-ai (Puinave), 556
tlalcacahoatl, 383
xochicacahoatl, 383
tlapal, 408
xocoatl, 379
Tlapalcacauatl, 408
too-soo (Yauna), 566 yagabizoya (Reko), 512
Toxoptera aurantii, 408, 432 yurac-cacao, 566, 570
V, 5. GOVERNMENT PRINTING OFFICE*

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CACAO AND ITS ALLIES

A TAXONOMIC REVISION OF THE GENUS THEOBROMA

"Celebrem etiam per universam Americam multique usua

fructum Cacao appellatum."

"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

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.

basin. I was fascinated by the unique structure of the flowers of the

cocoa tree, and its extraordinary fruit. My explorations from 1942

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

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,

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

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

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

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

as different others which are actually synonyms. Twenty-two species

are recognized in the present work. The number of species in the