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Taxonomy and classification of cacao

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DOI: 10.19103/AS.2017.0021.01

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BURLEIGH DODDS SERIES IN AGRICULTURAL SCIENCE

Achieving sustainable
cultivation of cocoa
Edited by Professor Pathmanathan Umaharan
Cocoa Research Centre - The University of the West Indies,
Trinidad and Tobago

E-CHAPTER FROM THIS BOOK

Taxonomy and classification of cacao
Ranjana Bhattacharjee, International Institute of Tropical Agriculture (IITA), Nigeria; and
Malachy Akoroda, Cocoa Research Institute of Nigeria, Nigeria

1 Introduction
2 Taxonomy and classification
3 The three groups of cocoa varieties
4 Conclusions
5 Where to look for further information
6 References

1 Introduction
Cacao is one of the economically important tree crops in West and Central Africa and is
native to the tropical forests of South America. The botanical name of cacao is Theobroma
cacao, first defined by Carolus Linnaeus, the father of modern-day taxonomic plant
classification, and was published in his classic book, Systema Naturae in the mid-1700s.
Theobroma is a Greek word that can be translated as ‘Food of Gods’: ‘theos’ meaning
‘god’ and ‘broma’ meaning ‘food’ (Wikipedia, 2013). The word ‘cacao’ is derived from the
Olmec and the subsequent Mayan languages (Kakaw), and the word ‘chocolate’ is derived
from the Nahuatl (the Aztec language) term ‘cacahuatl’, which is in turn derived from
Olmec/Mayan etymology (Dillinger et al., 2000). The other common names are cocoa
(English), cacaoyer (French), cacaoeiro/cacau (Portuguese/Brazil), kakao (Swedish), árbol
del cacao/cacaotero/calabacillo/forester (Spanish) and echter Kakaobaum (German). There
is archaeological evidence that the Mayans cultivated cacao 2000–4000 years before the
Spanish arrived in South America (Bergmann, 1969; Purdy and Schmidt, 1996; Whitlock
et al., 2001; Henderson et al., 2007; Powis et al., 2011).

1.1 Distribution of cacao
Whitlock et al. (2001) suggested the region extending from the forests of the Amazon to
the Orinoco and Tabasco in Southern Mexico to be the centre of origin of cacao (Van Hall,
1914). Cheesman (1944) believed the Upper Amazon near the Colombian–Ecuadorian
border, on the eastern flanks of the Andes, to be its centre of origin. He argued that
although cacao cultivation has been practised in Mexico and Central America for over
2000 years, there was no evidence of its wild populations in the region. Later, Schultes

http://dx.doi.org/10.19103/AS.2017.0021.01
© Burleigh Dodds Science Publishing Limited, 2018. All rights reserved.

Chapter taken from: Umaharan, P. (ed.), Achieving sustainable cultivation of cocoa, Burleigh Dodds Science Publishing,
Cambridge, UK, 2018, (ISBN: 978 1 78676 168 2; www.bdspublishing.com)
2 Taxonomy and classification of cacao

(1984) hypothesised that cacao dispersed along two routes from the Amazon Valley, one
leading to the north and the other the west. In this way, the domestication of cacao was
thought to have occurred in South America and then spread to Central America and
Southern Mexico, carried by migrating Indians. It is also estimated that the Mayas and
Aztecs domesticated and consumed cocoa, while the Mayas, Olmecs, Toltecs and Aztecs
used cocoa beans as the bitter drink, chocolatl, as well as a currency for trading, at least
1400 years ago (Purdy and Schmidt, 1996; Rössner, 1997; Nair, 2010; Anon, 2011). Several
expeditions and studies have shown that cocoa was first cultivated in Mexico by the Aztecs
before it spread to the Caribbean islands. Later, Hernán Cortés, a Spaniard, carried cocoa
to Spain in 1520s and used it as a beverage as well as a crop for cultivation. It was the
Spaniards who also introduced the crop into Equatorial Guinea in the seventeenth century.

1.1.1 Subspecies and domesticated cacao

Cuatrecasas (1964) suggested the simultaneous origin of cacao separately in South and
Central America, a hypothesis supported by most subsequent authors (Cope, 1976;
Wood and Lass, 1985; Gómez-Pompa et al, 1990; Laurent et al, 1994; de la Cruz et al,
1995; Whitkus et al, 1998). He also proposed that North and South American cacao
populations developed into two forms, separated by the Panama Isthmus, and that
this can be attributed to the great morphological diversity observed in Central as well
as in South America. These populations evolved independently and are recognised as
separate subspecies (T. cacao ssp. cacao and T. cacao ssp. sphaerocarpum). Furthermore,
Cuatrecasas (1964) hypothesised that wild plants from the Lacandon rainforest from
Mexico may possibly be ancestors of domesticated cacao. The subspecies proposed
by Cuatrecasas correspond to the two morphogeographic groups namely Criollo, which
corresponds to T. cacao ssp. cacao, and Forastero, which corresponds to T. cacao ssp.
sphaerocarpum, proposed by Cheesman (1944). A third hybrid group was called Trinitario,
originating from crosses between Criollo and Forastero. The Forastero group is composed
of very diverse populations with different geographic origins such as Upper Amazon,
Lower Amazon, Orinoco and the Guianas and can be identified according to the pod
morphology, for example, the Amelonado type.

2 Taxonomy and classification

Theobroma is exclusively neotropical in origin and its natural dispersion is in the tropical
lowland rainforests which extend from the Amazon basin through Southern Mexico (18°N to
15°S) (Cuatrecasas, 1964). Several studies on its morphological, anatomical, palynological
and chemical characteristics as well as its DNA sequences have confirmed the inclusion of
the family Sterculiaceae in a broadly defined Malvaceae sensu latu (Judd and Manchester,
1997; Alverson et al., 1999; Bayer et al., 1999), tribe Byttnerieae (Bentham and Hooker,
1862) or Theobromeae (Hutchinson, 1967; Takhtajan, 1997). There is minimal evidence
on the practice of cacao cultivation by the various indigenous groups of the Amazonian
rainforest (Sánchez et al., 1989; Dias, 2001a,b; Bartley, 2005). The wild relatives of cacao
thus include two different groups of germplasm. The first is the large spectrum of wild
populations which grow simultaneously in the Amazonian rainforest, ranging from French

© Burleigh Dodds Science Publishing Limited, 2018. All rights reserved.

Taxonomy and classification of cacao 3

Guiana to Bolivia. There is no reproductive barrier between cultivated and wild cacao
trees. Morphological characteristics of even the ancient cultigens, Criollo cacao from
Mesoamerica, are similar to those of their wild counterparts (Iwaro et al., 2005). Wild cacao,
therefore, may be used in breeding or commercial production, either as progenitors or as
clones (Dias and Resende, 2001; Eskes and Efron, 2006). The second category of wild
cacao is that of the 22 related Theobroma species (Cuatrecasas, 1964), although these
show crossing barrier with T. cacao. Three scholarly publications to date have recognised
a different number of species of Theobroma before Cuatrecasas (1964), which included 18
species (Bernoulli, 1869), 11 species (Schumann, 1886) and 13 species (Chevalier, 1946).
Bernoulli (1869) described five categories based on floral morphology and fruit structure.
These are:
Cacao: Petal ligule stipitate; staminodes erect, subulate; stamens 2-antheriferous;
calyx 5-parted, the laciniae equal. Ovate-oblong fruit included four species: T. cacao L.;
T. pentagona, ‘cacao lagarto’ from Guatemala; T. leiocarpa, ‘cumacaco’ from Guatemala;
and T. saltzmanniana from Bahia, Brazil.
Oreanthes: Petal appendix subsessile; staminodes erect, subulate; stamens
3-antheriferous; calyx 5-parted, the laciniae equal, included three species: T. speciosa
Willd. Ex Spreng.; T. quinquenervia and T. spruceana, all from Brazil.
Rhytidocarpus: Petal appendix subsessile; staminodes erect, claviform; stamens
2-antheriferous; calyx 5-parted, the laciniae equal; fruit woody, included two species:
T. bicolour Humb. & Bonpl. (synonym, T. ovatifolia DC.) and T. glauca Karst.
Telmatocarpus: Petal appendix; staminodes erect, linear-subulate with broad base;
stamens 3-antheriferous; calyx 5-parted, the laciniae equal; fruit ovate, lacunose, included
only one species: T. microcarpa Mart.
Glossopetalum: Petal ligule stipitate; staminodes reflexed, petaloid; stamens
3-antheriferous; calyx irregularly 3-5-fid, foliaceous; fruit sublignose. This is the largest
section, including eight species: T. augustifolia DC.; T. subincana Mart.; T. sylvestris Mart.;
T. macrantha from Brazil; T. ferruginea from Peru; T. obovata from Brazil; T. alba from British
Guiana and T. nitida from Brazil.

2.1 Classification
Morris (1882) was the first to publish the nomenclature of the cultivars of Theobroma
cacao. He distinguished two classes, the second of which was divided into the following
several varieties:

•• Class I: Cacao Criollo (Red)

•• Class II: Cacao Forestero
∘∘ Var. a. Cundeamor verugoso Amarillo (rough yellow Cerasee)
∘∘ Var. b. Cundeamor verugoso Colorado (rough red Cerasee)
∘∘ Var. c. Liso amarillo (smooth yellow)
∘∘ Var. d. Liso colorado (smooth red)
∘∘ Var. e. Amelonado amarillo (yellow melon)
∘∘ Var. f. Amelonado colorado (red melon)
∘∘ Var. g. Calabacillo amarillo (yellow calabash)
∘∘ Var. h. Calabacillo colorado (red calabash)

© Burleigh Dodds Science Publishing Limited, 2018. All rights reserved.

4 Taxonomy and classification of cacao

In 1892, Hart modified Morris’ classification of the varieties of T. cacao and distinguished
the following classes:

•• Class I: Criollo
•• Trinidad Criollo
∘∘ var. a. Amarillo (yellow, thick-skinned, bottle-necked)
∘∘ var. b. Colorado (red, thin-skinned, bottle-necked)
•• Venezuelan Criollo
∘∘ var. a. Amarillo (yellow, thick-skinned, high-shouldered, sometimes pointed)
∘∘ var. b. Colorado (red, thick-skinned, high-shouldered, sometimes pointed)
•• Nicaraguan Criollo
∘∘ var. a. Amarillo (yellow, thick-skinned, high-shouldered, very large beans with
light-coloured interior)
∘∘ var. b. Colorado (red, thick-skinned, high-shouldered, very large beans with
light-coloured interior)

•• Class II: Forastero

∘∘ var. a. Cundeamor verugoso Amarillo (yellow-warted)
∘∘ var. b. Cundeamor verugoso Colorado (red-warted)
∘∘ var. c. Ordinary amarillo (yellow forastero)
∘∘ var. d. Ordinary colorado (red forastero)
∘∘ var. e. Amelonado amarillo (yellow, melon-shaped)
∘∘ var. f. Amelonado Colorado (red, melon-shaped)

•• Class III: Calabacillo

∘∘ var. a. Amarillo (yellow, flat-beaned, smooth, thin- or thick-skinned, small pods)
∘∘ var. b. Colorado (red, flat-beaned, smooth, thin- or thick-skinned, small pods)

Theobroma pentagona

Theobroma pentagona (alligator cacao: yellow, heavily warted pods with five distinctly
raised ribs, large beans having white- or light-coloured interior).

2.2 Diversity and varieties

In 1944, Cheesman concluded that the whole assemblage of wild, semi-wild and cultivated
cacao constitutes one interbreeding population and supported the main division of
cacao into two groups of varieties, Criollo and Forastero. He divided Criollo into Central
American and South American Criollos. The Forasteros were divided into Amazonian
Forasteros, which can be found wild in Amazonia, and which are widespread in cultivation,
and the Trinitarios, possibly originating from the mingling of South American Criollo and
Amazonian Forastero stocks. He suggested that T. pentagonum was a simple form of
T. cacao, probably a segregate of the large cross-fertilised population. Cuatrecasas (1964)
described 22 species based on the classical method of comparative morphology such
as the structure of fruit and vegetative characteristics. He classified the species into six
sections: Andropetalum (T. mammosum Cuatr. & León); Glossopetalum [T. angustifolium
Moçiño & Sessé, T. canumanense Pires & Fróes, T. chocoense Cuatr., T. cirmolinae

© Burleigh Dodds Science Publishing Limited, 2018. All rights reserved.

Taxonomy and classification of cacao 5

Cuatr., T. grandiflorum (Willd. ex Spreng.) Schum., T. hylaeum Cuatr., T. nemorale Cuatr.,

T. obovatum Klotzsch ex Bernoulli, T. simiarum Donn. Smith., T. sinuosum Pavón ex Hubber,
T. stipulatum Cuatr., T. subincanum Mart.]; Oreanthes (T. bernouillii Pittier, T. glaucum Karst.,
T. speciosum Willd., T. sylvestre Mart., T. velutinum Benoist); Rhytidocarpus (T. bicolor
Humb. & Bonpl.); Telmatocarpus (T. gileri Cuatr., T. microcarpum Mart.) and Theobroma
(T. cacao L.). Representative species from all the sections can be found in Brazil, except
for Andropetalum (T. mammosum). The species which occur in, and are restricted to, the
Amazon basin of Brazil are: T. grandiflorum, T. obovatum, T. subincanum, T. speciosum,
T. sylvestre, T. microcarpum, T. glaucum, T. canumanense, T. bicolor and T. cacao,
(Cuatrecasas 1964; Figueira et al., 1994). The existence of the species T. canumanense is
based on the description of two exsiccates collected near the river Canumão (municipality
of Borba, AM).

2.2.1 Scientific classification/Taxonomic tree

Preferred scientific name: Theobroma cacao L.
Preferred common name: Cocoa
Other scientific names: Theobroma pentagonum Bernoulli; Theobroma sativum (Aubl.)
Lign & Le Bey
Domain: Eukaryota
Kingdom: Plantae
Subkingdom: Tracheobionta (Vascular plants)
Superdivision: Spermatophyta (Seed plants)
Division: Magnoliophyta (Flowering plants)
Phylum: Spermatophyta
Subphylum: Angiospermae
Class: Magnoliopsida (Dicotyledons)
Subclass: Dilleniidae
Order: Malvales
Family: Malvaceae/Sterculiaceae
Genus: Theobroma
Species: Theobroma cacao

Previous phylogenetic studies (Whitlock and Baum, 1999; Silva and Figueira, 2005) have
indicated that Theobroma is related to Herrania Goudot, a genus of about 20 species
monographed by Schultes (1958), both representatives of the Theobromeae (Whitlock
et al., 2001) along with two other genera, Glossostemon Desf., with one species from Arabia,
and the Neotropical Guazuma Mill., with 2–5 species. The classification of Theobroma and
Herrania into sections was through two clades. The Herrania clade was subdivided into the
relatively plesiomorphic section Subcymbicalyx and the section Herrania, with apomorphic
characteristics. The Theobroma clade was subdivided into a group containing sections
Andropetalum and Glossopetalum, which exhibit many plesiomorphic characteristics,
and a second group of the morphologically derived sections Oreanthes, Rhytidocarpus,
Telmatocarpus and Theobroma (Whitlock and Baum, 1999; Silva and Figueira, 2005).
Bernoulli (1869) quoted Herrania as a genus differing from Theobroma in the habit of the
plant and with respect to 5-6-foliolate digitate leaves. However, Theobroma and Herrani
species are both diploid with 2n = 20 chromosomes (Muñoz Ortega, 1948).

© Burleigh Dodds Science Publishing Limited, 2018. All rights reserved.

6 Taxonomy and classification of cacao

2.2.2 Effects of domestication
It is now clear that cacao was brought to Mesoamerica by early humans where its use and
domestication may have started around 4000 years ago (Powis et al., 2011). Humans are
also responsible for distributions of this species over the Amazon Basin (Clement et al.,
2010). In the case of both the Theobroma and Herrania species, the use of the sweet
pulp surrounding the seeds for making a fermented drink is the probable reason for its
domestication during the time of European contact (Patiño Rodriguez, 2002), a practice
which persists even today (Thomas et al., 2012). Recent discoveries in Honduras showed
that the Olmec people in the Ulua valley made an alcoholic drink in this way (Henderson
et al., 2007). The diverse nature of cacao’s uses as well as its use as a currency led to its
wide distribution in Mesoamerica before the Spaniards arrived (Bergmann, 1969; Young,
1994). However, only a very small portion of its diversity has been spread from Amazon to
Mesoamerica, and therefore domesticated cacao has a very narrow genetic background
(Dias et al., 2001; Bartley, 2005). The order Malvales is large and contains the family
Malvaceae to which cacao botanically belongs to.

2.2.3 Species in Theobroma
Infra.
Name Status Confidence level Source Date supplied rec.
Theobroma angustifolium Accepted TRO 18 April 2012
Sessé & Moc. ex DC.
Theobroma balaense Unresolved WCSP (in 26 March 2012
(P.Preuss) De Wild. review)
Theobroma bernoullii Accepted TRO 18 April 2012 2
Pittier
Theobroma bicolor Accepted WCSP (in 26 March 2012
Humb. & Bonpl. review)
Theobroma cacao L. Accepted WCSP (in 26 March 2012 1
review)
Theobroma calodesme Unresolved WCSP (in 26 March 2012
Diels review)
Theobroma canumanense Accepted TRO 18 April 2012
Pires & Fróes ex Cuatrec.
Theobroma caribaeum Unresolved WCSP (in 26 March 2012
Sweet review)
Theobroma celtifolia Unresolved TRO 18 April 2012
Salisb.
Theobroma cirmolinae Accepted TRO 18 April 2012
Cuatrec.
Theobroma duckei Huber Accepted TRO 18 April 2012

Theobroma ferrugineum Unresolved WCSP (in 26 March 2012

Bernoulli review)

© Burleigh Dodds Science Publishing Limited, 2018. All rights reserved.

Taxonomy and classification of cacao 7

Theobroma fossilium E.W. Unresolved TRO 18 April 2012

Berry
Theobroma gileri Cuatrec. Accepted TRO 18 April 2012

Theobroma glaucum H. Accepted TRO 18 April 2012

Karst.
Theobroma grandiflo- Accepted WCSP (in 26 March 2012
rum (Willd. ex Spreng.) review)
K.Schum.
Theobroma guyanense Unresolved WCSP (in 26 March 2012
Voigt review)
Theobroma hylaeum Accepted TRO 18 April 2012
Cuatrec.
Theobroma macranthum Unresolved WCSP (in 26 March 2012
Bernoulli review)
Theobroma mammosum Accepted TRO 18 April 2012
Cuatrec. & J. León
Theobroma martianum Unresolved WCSP (in 26 March 2012
D.Dietr. review)
Theobroma microcarpum Accepted WCSP (in 26 March 2012
Mart. review)
Theobroma montanum Unresolved WCSP (in 26 March 2012
Goudot ex Bernoulli review)
Theobroma nemorale Accepted TRO 18 April 2012
Cuatrec.
Theobroma obovatum Accepted WCSP (in 26 March 2012
Klotzsch ex Bernoulli review)
Theobroma ovatifolium Unresolved WCSP (in 26 March 2012
DC. review)
Theobroma pulcherrimum Unresolved WCSP (in 26 March 2012
Pittier review)
Theobroma quinquen- Unresolved WCSP (in 26 March 2012
ervium Bernoulli review)
Theobroma sagittatum Unresolved WCSP (in 26 March 2012
Pav. ex A.Chev. review)
Theobroma salzmanni- Unresolved WCSP (in 26 March 2012
anum Bernoulli review)
Theobroma simiarum Accepted TRO 18 April 2012
Donn. Sm.
Theobroma sinuosum Pav. Accepted TRO 18 April 2012
ex Huber
Theobroma speciosum Accepted WCSP (in 26 March 2012
Willd. ex Spreng. review)

© Burleigh Dodds Science Publishing Limited, 2018. All rights reserved.

8 Taxonomy and classification of cacao

Theobroma stipulatum Accepted TRO 18 April 2012

Cuatrec.
Theobroma subincanum Accepted WCSP (in 26 March 2012
Mart. review)
Theobroma sylvestre Accepted TRO 18 April 2012
Aubl. ex Mart. in Buchner
Theobroma velutinum Accepted TRO 18 April 2012
Benoist

Statistics of the Plant List includes 71 scientific plant names of species rank for the genus
Theobroma. Of these 22 are accepted species names.

Status Total
Accepted 25
Synonym 46
Unplaced 0
Unassessed 15

2.2.4 Disagreements on classification
The botanical taxonomy of cacao is not straightforward. The high level of synonyms is a
hindrance to good classification and there are aspects of disagreement as shown below:

Overall, the taxonomy of Theobroma cacao L. is as follows:

Kingdom: Plantae (plants)
Infrakingdom: Streptophyta (land plants)
Subkingdom: Tracheobionta (vascular plants)
Subkingdom: Viridiplantae
Superdivision: Embryophyta
Superdivision: Spermatophyta (seed plants)
Division: Tracheophyta (vascular plants, tracheophytes)
Division: Magnoliophyta (flowering plants)
Subdivision: Spermatophytina (spermatophytes, seed plants, phanérogames)
Class: Equisetopsida
Class: Magnoliopsida (dicotyledons, e.g. coffee)
Subclass: Dilleniidae (e.g. like tea)
Subclass: Magnoliidae
Superorder: Rosanae
Order: Malvales
Family: Malvaceae (formerly Sterculiaceae)
Genus: Theobroma
Other species of family Theobroma
T. grandiflorum
T. bicolor

© Burleigh Dodds Science Publishing Limited, 2018. All rights reserved.

Taxonomy and classification of cacao 9

Since the mid-twentieth century, T. cacao varieties have been classified into three general
groups: ‘Criollo’, ‘Forastero’ and ‘Trinitario’, based on morphological descriptors and
geographic origin (Engels, 1981; Enriquez and Soria, 1967). Recent genetic studies indicate
that this broad categorisation is overly simplistic with regard to the term ‘Forastero’ and
fails to acknowledge the significant amount of genetic variation contained within this
group. In contrast, ‘Criollo’ refers to a group of genetically similar trees which produce
lightly pigmented seeds and share several other morphological traits. ‘Trinitario’ refers to
hybrids of the ‘Criollo’ and ‘Forastero’ strains.

3 The three groups of cocoa varieties

3.1 Criollo
The Criollo group refers to genetically similar trees, believed to have been domesticated
by the Mayan civilisation. The seeds (beans) tend to be round in profile and contain white
or pale purple cotyledons. Young seedlings can be identified by the presence of green
cotyledons and by that of leaf stems (petioles) with a horizontally opposed orientation.
This variety displays low vigour, poor productivity and high susceptibility to diseases,
insects and stress and is therefore less widely cultivated (Afoakwa, 2010). Until the mid-
eighteenth century, Criollo was the most commonly cultivated type of cacao; however,
the vast majority of these trees have been replaced with more vigorous and hardy trees of
hybrid or ‘Forastero’ ancestry. This type is very rare and is only found in old plantations in
Venezuela, Central America, Madagascar, Sri Lanka and Samoa (Fowler, 2009). The yield
of Criollo plantations is lower than that of Forastero. Notable characteristics include a
unique growth habit that frequently lacks the usual whorl of five branches (jorquette) and
consists entirely of plagiotropic stems with occasional bi- and trifurcating branches. Fruits
typically have a soft thin husk with a textured surface and usually have some degree of red
pigmentation. Chocolate made from Criollo is light in colour and has a subtle or delicate
taste that is low in basic chocolate flavour. Criollo beans are often sold at higher prices,
which can somewhat offset their lower yield. Criollo may be translated as ‘native’ or ‘first
grown’ in Spanish-speaking countries of the Americas. It should be noted that so-called
Criollo cacao in many countries may not have the typical genetic profile of pure Criollo.

3.2 Forastero
Forastero means ‘foreigner’ in Spanish. It refers to any trees that are not Criollo or a
hybrid and which usually produce deep purple seeds (Hebbar et al., 2011). The term
encompasses a wide range of distinct populations with unique characteristics and is not a
meaningful descriptive term. Forastero is native to the Amazon region and largely grown
in West Africa and Southeast Asia. It forms 95% of world cocoa production (Afoakwa,
2010; Fowler, 2009; Afoakwa et al., 2012) and is the most widely used due to its higher
yield than the Criollo variety. Because of the high genetic variability within this group,
Forastero types exhibit greater variability in tree and fruit morphology and are generally
more vigorous and less susceptible to disease and pests than are Criollo trees. When
ripe, the pods are hard and yellow, have a rounded melon-like shape and contain 30 or
more beans ranging in colour from pale to deep purple. Several landraces are particularly
well known. Grown mainly in Brazil and Africa, it is hardier, higher yielding and easier

© Burleigh Dodds Science Publishing Limited, 2018. All rights reserved.

10 Taxonomy and classification of cacao

to cultivate than Criollo and is used in every blend of chocolate that is produced. The
Amelonado type is widely grown throughout Western Africa and produces the majority
of ‘bulk cacao’ beans. Although Forastero beans are said to produce chocolate rich in
chocolate flavour but low in complex or fruit flavour notes, there are several well-known
exceptions. Forastero may be subdivided into Upper Amazonian [wild or semi-wild
cacaos as described by Pound (1938)] and Lower Amazonian, characterised by a rather
uniform pod type called Amelonado (non-pigmented, smooth, melon-shaped pods with
a blunt end). Lower Amazonian cacaos now constitute the most prevalent cultivated type
worldwide and are grown notably in West Africa and Brazil. These types grow wild in the
Guyanas and the eastern Brazilian Amazon and may have been partially domesticated by
pre-Colombian Amazonian natives for their aromatic pulp (Barrau, 1979).

3.3 Trinitario and other hybrids

In the strict sense, Trinitario refers to clones, or the progenies, of hybrids originally produced
in Trinidad between ‘Criollo’ and ‘Forastero’ trees which originated in the lower Amazon
River basin (Willson, 1999). However, it later spread to Venezuela, Ecuador, Cameroon,
Samoa, Sri Lanka, Java and Papua New Guinea (ICCO, 2012). It is believed that it first
came into existence on the island of Trinidad, after a hurricane nearly destroyed the local
Criollo crops in 1727. Some Trinitario varieties produce cocoa beans with special flavours.
They have hard pods and are variable in colour, containing 30 or more beans of variable
colour, although white beans are rare. Trinitario is considered to be either an intermediate
type between Criollo and Forastero (Lockwood and Gyamfi, 1979) or a group of hybrids
displaying characteristics which include the total range of variation (Cheesman, 1944). The
Trinitarios combine the best characteristics of the two main varieties: the hardiness and
high yield of Forestero and the refined taste of Criollo. Trinitario types have not been found
under wild conditions. Because Criollos and Forastero types are so heterogeneous, the
resulting hybrids might not be distinct from the parental populations, making Trinitarios
impossible to define except by geographical origin (B.G.D. Bartley, unpublished).

3.4 Nacional
The Nacional variety grows in Ecuador and is believed to have originated from the
Amazonian area of Ecuador (Fowler, 2009; Afoakwa, 2010). It has distinct characteristics of
aroma and flavour. Although less widely cultivated and contributing to only 5% of global
cocoa production, it represents more than 50% of the fine cocoa marketed worldwide
every year. Until the beginning of twentieth century, Nacional cultivars were the only type
of cocoa cultivated in Ecuador (Soria, 1966), with the unique flavour and aroma, known
as ‘arriba’. The fine flavoured quality chocolate products obtained from Nacional cocoa
beans are highly valued by chocolate manufacturers. However, the original Nacional
cocoa trees are currently in danger of extinction due to the introduction of an external
unrelated germplasm (Solorzano et al., 2012). A large genetic admixture between native
Nacional cocoa and foreign germplasm is currently found in modern Ecuador plantations
(Solorzano et al., 2009), thus reducing the fine-flavoured cocoa aroma. Currently, pure
Nacional cocoa varieties are rare and there is an increasing demand for fine flavour cocoa
although Ecuador still represents 6.8% of the supply of ‘arriba’ flavoured cocoa in the
world market (ICCO, 2012). Silva et al. (2004) provided a key to the species and hybrids
of genus Theobroma.

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Taxonomy and classification of cacao 11

3.5 Proposed new classification

Motamayor et al. (2008) proposed a new classification into ten major groups within the
Forastero cacao, based on genetic diversity in varieties from Latin America. These are
Marañon, Curaray, Criollo, Iquitos, Nanay, Contamana, Amelonado, Purús, Nacional and
Guiana. This new classification more accurately reflects the genetic diversity of cocoa (Fig. 1).
Since 2008, several scientific studies have been carried out to assess the genetic diversity
of cacao (Zhang et al., 2009, 2011; Trognitz et al., 2011; Ji et al., 2013; Santos et al., 2015).
However, most of these are limited to only a few geographical locations and to three or
four groups (Criollos, Foreseteros, Trinitariors and or Nacional or Cacao Nacional Boliviano
types). Thomas et al. (2012) used spatial diversity patterns to explain genetic differentiation
in cacao through the possible contribution of climate change, geographical locations,
history and culture. However, there are several questions which remain unanswered and
a comprehensive study combining application of molecular and modelling approaches is
needed to understand the entire genetic diversity and population structure of cacao.

Figure 1 Neighbour joining tree from Cavalli-Sforza and Edwards genetic distance [16] matrix among
the 36 subclusters identified using STRUCTURE (559 clones). Source Motamayor et al. (2008).

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12 Taxonomy and classification of cacao

Figure 2 Species richness of the genus Theobroma. Source Thomas et al. (2012).

4 Conclusions
It has been concluded by many authors that the centre of genetic diversity of cocoa is
located in the upper Amazonian regions bordering Peru, Brazil, Colombia and Ecuador,
and to a lesser extent, in the southern part of Central America (Motamayor, et al., 2002;
Sereno et al., 2006; Zhang, et al., 2006).
A certain level of overlap exists between the areas of highest species richness of
the genus Theobroma and the area where the highest genetic diversity of cacao is
observed. This corroborates the hypothesis that cacao also originated in the Western
Upper Amazon. Interpretation of the Pleistocene modelled distribution of cacao,
together with present-day genetic observations, suggests that the species was already
distributed widely across the Western Amazon. The current distribution of cocoa is likely
to have been partly shaped by varying degrees of human interventions. The fact that
the pulp of several other species in the genus Theobroma was used in a similar fashion
as cacao by South-American indigenous groups (Patiño Rodriguez, 2002) might explain
why the present-day highest species richness of the genus Theobroma is located in this
area (Fig. 2).

5 Where to look for further information

The demand for high-quality cocoa is increasing which necessitates the need to broaden
the genetic diversity by looking at additional cocoa germplasm from the regions that has
not yet been explored. A lot of germplasm characterization research is being undertaken
at USDA-ARS, Beltsville in collaboration with the international gene bank, Trinidad and
Tobago. Future research should therefore concentrate on the development of cocoa
varieties that can give higher yields and produce quality beans in the light of changing
climate and associated threats of biotic and abiotic stresses. Although a lot of work and
publications are available on defining the origin, taxonomy and classification of cacao, the
questions still remains if there are other members of the genus Theobroma that produces
cocoa-like products and lot of information can be available in ICCO, WCF and CABI
websites.

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Taxonomy and classification of cacao 13

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