The ten genetic groups of cacao

Bars and books still divide cacao into Criollo, Forastero and Trinitario. Geneticists stopped in 2008, when a study of more than a thousand trees found ten groups instead. These are the ten.

10 groups952 trees analysed3 papers read in full

The short version

In 2008 Juan Carlos Motamayor and colleagues genotyped 1,241 cacao trees from germplasm collections and proposed sorting the species into ten groups, named after a place or after a traditional cultivar: Marañón, Curaray, Criollo, Iquitos, Nanay, Contamana, Amelonado, Purús, Nacional and Guiana. [1] Ten years later, sequencing 200 whole genomes found the same ten. [2]

The practical consequence is that Forastero is not a kind of cacao. It was the word for everything that was not Criollo, and it covers nine populations that are strongly differentiated from one another.

The ten groups

In the order the paper lists them. Where an entry says little, that is all the papers read for this page say about it in their text.

Marañón

Amazonian populationPA clones

Named for: The Marañón river in Peru, the place most represented among its trees.

  • Trees of the PA series, from F. J. Pound, whose upper-Amazon collection is the most widely used in breeding, make up 68% of the individuals the paper placed in this group. [1]

Curaray

Amazonian population

Named for: The Curaray river; a later paper describes the group as from northern Ecuador and southern Colombia.

  • Whole-genome analysis found this to be the population most closely related to Criollo, and the authors suggest domesticated cacao began as a subset of ancient Curaray trees. [2]

Criollo

Traditional cultivar

Named for: The traditional cultivar of Central America and Mexico, whose name the classification kept.

  • The only group found in Central American forests, where the others occur only on farms. All ten groups, Criollo included, are represented in South America, which the authors take as evidence against wild cacao having evolved in Central America. They add a caution: what now looks like primary forest there may have been cultivated before Columbus. [1]
  • So uniform that trees collected in different places and grown from seed were identical at all 96 genetic markers used. [1]
  • Genome sequencing dated its domestication to about 3,600 years ago, with a very wide margin (95% interval 2,481 to 13,806 years), from a small founding population and as a single event. [2]
  • It carries a higher proportion of harmful mutations than the other groups, and in a set of trees with measured yields, yield fell as the share of Criollo ancestry rose. [2]

Criollo as a variety, in the catalogue

Iquitos

Amazonian populationIMC clones

Named for: The city of Iquitos on the Peruvian Amazon, where several major rivers meet.

  • Trees from several different groups have been collected around Iquitos itself, which the authors think may reflect ancient or modern movement of cacao by people along the rivers. [1]

Nanay

Amazonian populationNA clones

Named for: The Nanay river, near Iquitos.

  • Nine in ten of the individuals in this group come from one collecting expedition, and the group is so narrow genetically that separately collected trees could not be told apart with 96 markers. [1]

Contamana

Amazonian populationSCA clones

Named for: Contamana, on the Ucayali river in Peru.

  • The Scavina clones belong here. The collector's report was read as meaning they descend from a single tree, but SCA-6 and SCA-12 do not share a haplotype and so cannot both come from one mother tree. [1]

Amelonado

Traditional cultivar

Named for: The traditional cultivar of that name, grown in Bahia in Brazil and known in Ghana as West African Amelonado.

  • The group brings together wild trees from widely separated places with cultivated ones from Bahia in Brazil, Costa Rica and Ghana. The authors call their findings on its origin less clear than for Nacional, noting that it includes trees from the Pará river, where historical accounts place its domestication. [1]
  • Ancestry from Amelonado, Criollo and Nacional is over-represented in most of the mixed and bred trees analysed, and the other seven groups are correspondingly little used in breeding. [2]

Amelonado as a variety, in the catalogue

Purús

Amazonian population

Named for: The Purús river of the south-western Amazon.

  • Its trees from the upper Solimões and Içá rivers did not sit cleanly in any group, falling between this one and its nearest neighbour, which the authors attribute to cacao genes moving along the main river. [1]
  • The one possible exception to Criollo's isolation: the genome study found no later contribution to Criollo from any group except a potential recent one from Purús. [2]

Nacional

Traditional cultivarMO clones

Named for: The traditional cultivar of coastal Ecuador.

  • Although Nacional is grown on the Pacific side of the Andes, its group includes wild trees from the Amazonian side, along the Morona, Nangaritza and Zamora rivers. [1]
  • A later study with 169 wild and cultivated trees identified three wild trees from the Yacuambi and Nangaritza rivers as the most likely parents of the Nacional type, and proposed Zamora Chinchipe province in southern Amazonian Ecuador as where it was domesticated. [3]
  • Modern Ecuadorian plantations are described as a broad genetic mixture of the native variety with introduced cacao, in which the characteristic aroma has declined. [3]

Nacional as a variety, in the catalogue

Guiana

Amazonian population

Named for: The Guianas; the paper's wild material included trees collected in French Guiana.

  • The study's table of clones places 59 trees here: 56 collected in French Guiana, along the Camopi and neighbouring rivers, and 3 in Brazil. [1]

What became of the old names

Criollo
Kept as the name of one of the ten groups. It was always a traditional cultivar, and the genetics agree that it is one distinct, very uniform population. [1]
Forastero
Not a group. It meant everything that was not Criollo, and covers what are now nine separate, strongly differentiated populations. The paper proposes its ten groups in place of the old three names. [1]
Trinitario
Not a group either, but hybrids: of Criollo with Amelonado, and of those hybrids with other Amazonian trees. The 2008 study excluded them on purpose, as mixtures made by people. [1]
Nacional and Amelonado
Two traditional cultivars that the old scheme filed under Forastero. Each turned out to be a group of its own, and both names were kept. [1]

Trinitario’s parentage was established separately: a study of Trinitario trees from 17 countries concluded that they result from hybridisation between Criollo and Amelonado. [7] The catalogue’s own entries for Forastero and Trinitario treat them as the trade names they are.

Five clone codes, and where they sit

Clone names such as SCA-6 or SCA-12 begin with letters that say where a tree was collected. Five of the best-known series come from a single upper-Amazon collection, made by F. J. Pound, and the paper places each in a group. [1]

Clone series from the Pound collection, the group each belongs to, and the share of that group’s analysed trees they make up
CodeStands forGroupShare of the group’s trees
PAParinariMarañón68%
IMCIquitos MixedIquitos54%
NANanayNanay90%
SCAScavinaContamana28%
MOMoronaNacional27%

The last column matters more than it looks. Nine in ten of the Nanay trees in the study, and two in three of the Marañón ones, descend from that one expedition, in which, on one reading of the collector’s report, the seed for each series came from somewhere between one and twenty mother trees. The paper notes that this collection is the material most used by breeding programmes worldwide, so much of what breeders know as “upper Amazon” cacao is a narrow sample of it.

For how clone names work in general, see how to read cacao clone codes. Any individual clone in the study can be looked up below.

Look up a clone

The study published the full list of the 952 trees it kept, with the group each was placed in. The article is in the public domain, so the list is here to search and to download as a CSV file.

A match is a match on the name. The study set aside 23% of the trees it sampled because their labels could not be trusted, so a tree sold under a name in the table is not thereby the tree that was genotyped.

The table, group by group

For each genetic group: trees in the study’s table, how many were clearly assigned, where they were collected, and the named subgroups
GroupTreesClearly assignedCollected inNamed subgroups
Nanay152121Peru 152Nanay I; Nanay III; Nanay II
Marañón143130Peru 98, Brazil 45Ji-Paraná R; Parinari IV; Parinari I; Parinari II; Parinari III
Curaray11786Ecuador 116, Colombia 1Coca R - Napo R; Curaray R - Napo R; Upano R; Bobonaza R
Iquitos11773Peru 82, Brazil 27, Ecuador 7, Colombia 1IMC I; IMC II; Amazonas; Middle Solimões R
Purús11074Brazil 87, Ecuador 16, Colombia 7Acre R; Caeté R - Purús R; Upper Solimões R - Iça R; Purús R
Amelonado9463Brazil 79, French Guiana 8, Ecuador 3, Ghana 2, Colombia 1, Costa Rica 1Amelonado Type II; Amelonado Type I; Uatama R; Amelonado Type III
Contamana6959Peru 61, Brazil 7, Ecuador 1Scavina; Nanay VII; Vilcanota; Embira R
Guiana5951French Guiana 56, Brazil 3Camopi R-Euleupousing R; Camopi R; Kérindioutou R
Nacional5236Ecuador 28, Peru 23, Colombia 1Nacional; Morona
Criollo3939Mexico 17, Colombia 8, Venezuela 5, Panama 3, Belize 2, Nicaragua 2, Costa Rica 1, Ecuador 1Lacandona; Santa Marta; Lacadona; Belize

Counted from the table by this site’s build. “Clearly assigned” means a membership of 0.70 or more, the study’s own threshold; the rows total 732, three fewer than the paper’s text gives, which rounding in the displayed figures would explain. Subgroup names are as printed, including two spellings of one Criollo subgroup. The counts show how many trees were sampled, not how common a group is in the wild or on farms.

Not in the table: CCN-51

The clone most often asked about is absent, because the study left out hybrids. What is on record about its parentage comes from a peer-reviewed review.

  • CCN-51 is traditionally recorded as a three-way cross, (IMC-67 × ICS-95) × a tree known as Canelo or Oriente 1, made by Homero Castro Zurita on his farm in the Naranjal area of Guayas province, Ecuador, in response to witches' broom disease. [6]
  • The review reports a 2014 genotyping study by Boza and colleagues as confirming IMC 67 and ICS 95 as parents, with IMC 67 the larger contributor. The third parent, Oriente 1, was not found in any germplasm collection, so nothing genetic could be tied to it. [6]
  • The same work is reported as finding CCN-51 highly heterozygous and almost unrelated to Nacional: only about 1% of the alleles analysed were associated with Nacional ones. [6]

The catalogue’s entry is CCN-51.

How the ten were arrived at

  • 1,241 trees held in germplasm collections, originally gathered in 12 countries between 1937 and 2005, were typed at 106 genetic markers, of which 96 were kept.
  • 289 of them, 23%, were set aside before analysis as mislabelled, duplicated or hybrid. 100 pairs of supposedly different trees were genetically identical. The paper also cites an earlier estimate, made from collection records, that between 15% and 44% of trees in cacao germplasm collections are misidentified.
  • Of the 952 that remained, 416 came from Peru, 248 from Brazil and 172 from Ecuador.
  • 735 belonged clearly to one group. The rest were mixtures, and more genetically varied, which the authors read as trees from zones where populations meet.
  • Differences between the ten groups accounted for 38.1% of all the genetic variation measured, and the groups divide further into 36 subgroups.

The greatest diversity was in the upper Amazon of Peru and Brazil, in keeping with the species having originated there. The genome study later found diversity falling steadily from the Pacific side of the basin to the Atlantic side. [1] [2]

Where the trees are kept

The trees in these studies came from living collections: fields of cacao trees, each propagated from a named original. The studies were possible because those collections exist, and their limits are the collections’ limits.

  • More than 40 collections around the world hold over 24,370 accessions of cacao between them, with a good deal of duplication within and between collections. [4]
  • Two are international collections: the Cocoa Research Centre of the University of the West Indies in Trinidad, and CATIE in Costa Rica. Both have agreements under the international plant treaty to keep their collections for the long term and to supply material freely to any qualified institution or person. [4]
  • Outside those two there is little movement of cacao between countries. Plants that do travel go through quarantine, chiefly at the International Cocoa Quarantine Centre at the University of Reading, where they are tested for viruses. [4]
  • The Trinidad genebank was begun in 1982 on a site of about 100 acres at Centeno, by bringing together earlier collections: the 100 Imperial College Selections, the ICS clones, from F. J. Pound's survey of Trinidad and Tobago in the 1930s; selections from Marper Farm, which Pound set up after his upper-Amazon expeditions of 1937 to 1942; and other national collections. By 1994 over 2,000 different types had been planted. [5]
  • The network lists 32 institutes: the two international collections, the quarantine centre, and 29 national collections in 21 countries and territories, from Brazil and Peru to Ghana, Indonesia and Papua New Guinea. [4]
  • Mislabelling is acknowledged by the network itself, which puts misidentification of trees within collections at as high as 30%. Few collections keep a safety duplicate of their unique material at a second site. [4]
  • Although scientific breeding of cacao began more than 70 years before the strategy was written, only about a quarter of cacao farms were then planted with improved varieties. [4]

The counts are from a strategy written in 2012 and will have changed. The database that catalogues individual accessions, the International Cocoa Germplasm Database, could not be reached when this page was written, so no collection’s holdings are listed.

What this does not tell you

Nothing about flavour. None of these studies tasted anything by group, and a group name on a wrapper is not a flavour description.

Nothing certain about the tree behind a particular bar. Most cultivated cacao is a mixture of groups, the study left hybrids out by design, and it set aside 23% of the trees it sampled as mislabelled, duplicated or hybrid before it began.

Ten is the number these samples supported, not a law. The authors report more than one possible arrangement at ten groups, and collecting has been sparse and uneven: often only a few trees with a wanted trait were taken from a place.

The 2008 study used 96 markers; the dates and the yield finding come from one later genome study. The domestication date in particular has a margin of several thousand years.

Where these figures come from

Publisher
Motamayor and colleagues, in PLoS ONE — Table S3, 'List of the 952 retained clones', supplementary to Motamayor et al. 2008, PLoS ONE 3(10): e3311
Release used
Published 1 October 2008; retrieved 8 October 2026.
Covers
Trees held in germplasm collections, originally collected in twelve countries of South and Central America and, for two clones, Ghana. Collected 1937 to 2005; genotyped for a study published in 2008.
Units
One row per sampled tree. Membership coefficients run from 0 to 1 and are the study's estimate of how fully a tree belongs to its group.
Licence
Creative Commons Public Domain declaration (as stated in the article)
How it is revised
A supplementary file to a published paper; not revised. Later studies have re-examined individual clones and the file does not reflect them.
Next check here
8 October 2027. None. Recheck only that the file is still published.

What was changed

  • The single worksheet was converted to CSV; the nine data columns were kept and the column counting clustering schemes was dropped.
  • Runs of spaces inside clone names were collapsed to one space. No name was otherwise altered, and the table's own spellings were kept.
  • The two trailing rows (a blank and a footnote) were removed, leaving 952 rows.
  • Subcluster entries of N/A were left empty. Entries the table marks with an asterisk were kept as printed; the table does not explain the mark.
  • Coefficients were written to at most three decimal places, as displayed in the table.

What these figures cannot tell you

It records where a 2008 study placed the trees it sampled. A tree sold or planted under the same name elsewhere may not be the same genotype: the study itself set aside 289 trees as mislabelled, duplicated or hybrid.

Twenty clone names appear more than once with different laboratory identifiers, because seedlings of one mother tree were sometimes kept under her name. They are separate rows.

Hybrids were excluded by design, so Trinitario clones and modern bred clones such as CCN-51 are not in it. Absence from the table says nothing about a clone.

A group is not a flavour. Nothing in the table describes how any clone tastes.

732 rows carry a membership of 0.70 or more as displayed; the paper's text counts 735, a difference that rounding of the displayed figures would explain.

Download

Reusing them: Motamayor JC, Lachenaud P, da Silva e Mota JW, Loor R, Kuhn DN, Brown JS, Schnell RJ (2008). Geographic and genetic population differentiation of the Amazonian chocolate tree (Theobroma cacao L). PLoS ONE 3(10): e3311, Table S3. The article is published under a Creative Commons Public Domain declaration. Counts by group are ChocolateHQ calculations from the table.

Sources

  1. Motamayor JC, Lachenaud P, da Silva e Mota JW, et al. Geographic and genetic population differentiation of the Amazonian chocolate tree (Theobroma cacao L). PLoS ONE. 2008;3(10):e3311. doi.org/10.1371/journal.pone.0003311 (read: full text)
  2. Cornejo OE, Yee M-C, Dominguez V, et al. Population genomic analyses of the chocolate tree, Theobroma cacao L., provide insights into its domestication process. Commun Biol. 2018;1:167. doi.org/10.1038/s42003-018-0168-6 (read: full text)
  3. Loor Solorzano RG, Fouet O, Lemainque A, et al. Insight into the wild origin, migration and domestication history of the fine flavour Nacional Theobroma cacao L. variety from Ecuador. PLoS ONE. 2012;7(11):e48438. doi.org/10.1371/journal.pone.0048438 (read: full text)
  4. CacaoNet. A Global Strategy for the Conservation and Use of Cacao Genetic Resources, as the Foundation for a Sustainable Cocoa Economy. 2012. Web edition, section 'Where we are today', and the network's list of collections. www.cacaonet.org/global-strategy/where-we-are-today (read: web page)
  5. Cocoa Research Centre, The University of the West Indies. The International Cocoa Genebank, Trinidad. sta.uwi.edu/cru/international-cocoa-genebank (read: web page)
  6. Jaimez RE, Barragan L, Fernández-Niño M, et al. Theobroma cacao L. cultivar CCN 51: a comprehensive review on origin, genetics, sensory properties, production dynamics, and physiological aspects. PeerJ. 2022;10:e12676. doi.org/10.7717/peerj.12676 (read: full text)
  7. Motamayor JC, Risterucci AM, Heath M, Lanaud C. Cacao domestication II: progenitor germplasm of the Trinitario cacao cultivar. Heredity. 2003;91:322–330. doi.org/10.1038/sj.hdy.6800298 (read: abstract)

Reviewed 8 October 2026. The studies themselves are summarised one by one in cacao genetics research.