**2. Cacao introduction history and genetic materials**

Traditionally, cacao types cultivated are subdivided into three major '*geneti*c' groups: Forastero, Criollo (domesticated by the Amerindians in Central America), and Trinitario (hybrids between Forastero and Criollo, originating from Trinidad). While the Forastero trees are vigorous and more resistant to diseases, the Criollo trees are poor yielding and highly susceptible, although Criollo trees produce high premium quality beans with aromatic flavor. The Brazilian cacao of the Amelonado type (Lower Amazon Forastero) was first introduced by the Portuguese into Principe around 1822, and reached Sao Tomé in the 1850s (Bartley 2005). According to Nosti, quoted by Toxopeus (1964), it was from this collection that the Spaniards brought cacao into the Island of Fernando Po (now Bioko), Equatorial Guinea in 1854. This collection in Fernando Po became the major source of cacao introduced into mainland West Africa at several times by many persons including traders and migrant workers, agencies, missionaries among others. Available records showed that cacao was introduced from Fernando Po by workers and traders like Squiss Ibaningo into Nigeria in 1874, Tetteh Quarshie into Ghana in 1878, and Cote d'Ivoire in 1879 (Edwin and Masters 2005; N'Goran et al. 1992; Opeke 1969). Missionaries like the Basel missionaries, Royal Botanical Garden curators, colonial administrations played significant roles in the introduction of cacao types from different origins into the mainland West and Central Africa. These earlier introductions from Fernando Po formed the initial basis of cacao grown in West Africa, and was referred to as the "West African Amelonado". During the late nineteenth century, the Colonial administration also introduced some red-podded cacao materials from British West Indies into botanical gardens established in Aburi (Ghana) and Lagos (Nigeria) (Toxopeus 1964). By 1910, Ghana, followed by Nigeria, had become one of the largest producing countries, thus making the West Africa sub-region an important growing area critical to the sustainability of the world's cocoa economy, a status it still maintains today.

The introduction of cacao germplasm into island and mainland of Africa took place in response to two main waves of idea which naturally divided cacao germplasm introduction into: 1. Exploratory Colonial Period (1822 – 1909), and, 2. Expansionary Experimental Preand Post-Independence Period (1910 – 2010). During the first era which spanned early 19th to the end of the 19th century, cocoa seeds and plants were transported in barrels and shipments across the sea from the northern parts of Southern America and Central America to Africa. This was in response to the then imbibed and appreciated Aztec and Mayan culture of drinking '*chocolatl*', 'The Food of the Gods". The favourable similar tropical humid climatic conditions and abundant rainforest vegetation provided impetus for this transatlantic exploratory introduction which fortunately gave good results. The cultivation

cocoa is produced primarily in central southern Cameroon by millions of small scale farmers. In Cameroon, cocoa cultivation is currently one of the major sources of revenues of rural households (1 to 2 millions of people) of the forest agro-ecological zones in the country (South and South-Western parts). Cocoa is grown in more than 200 000 farms and the total cocoa growing surface is estimated to be 400 000 hectares (Efombagn et. al., 2006). In other countries such as Togo, Sierra Leone, Liberia, Equatorial Guinea, Sao Tome and Principe, Gabon and Democratic Republic of Congo, cocoa production makes significant agricultural contribution

Traditionally, cacao types cultivated are subdivided into three major '*geneti*c' groups: Forastero, Criollo (domesticated by the Amerindians in Central America), and Trinitario (hybrids between Forastero and Criollo, originating from Trinidad). While the Forastero trees are vigorous and more resistant to diseases, the Criollo trees are poor yielding and highly susceptible, although Criollo trees produce high premium quality beans with aromatic flavor. The Brazilian cacao of the Amelonado type (Lower Amazon Forastero) was first introduced by the Portuguese into Principe around 1822, and reached Sao Tomé in the 1850s (Bartley 2005). According to Nosti, quoted by Toxopeus (1964), it was from this collection that the Spaniards brought cacao into the Island of Fernando Po (now Bioko), Equatorial Guinea in 1854. This collection in Fernando Po became the major source of cacao introduced into mainland West Africa at several times by many persons including traders and migrant workers, agencies, missionaries among others. Available records showed that cacao was introduced from Fernando Po by workers and traders like Squiss Ibaningo into Nigeria in 1874, Tetteh Quarshie into Ghana in 1878, and Cote d'Ivoire in 1879 (Edwin and Masters 2005; N'Goran et al. 1992; Opeke 1969). Missionaries like the Basel missionaries, Royal Botanical Garden curators, colonial administrations played significant roles in the introduction of cacao types from different origins into the mainland West and Central Africa. These earlier introductions from Fernando Po formed the initial basis of cacao grown in West Africa, and was referred to as the "West African Amelonado". During the late nineteenth century, the Colonial administration also introduced some red-podded cacao materials from British West Indies into botanical gardens established in Aburi (Ghana) and Lagos (Nigeria) (Toxopeus 1964). By 1910, Ghana, followed by Nigeria, had become one of the largest producing countries, thus making the West Africa sub-region an important growing area critical to the sustainability of the world's cocoa

The introduction of cacao germplasm into island and mainland of Africa took place in response to two main waves of idea which naturally divided cacao germplasm introduction into: 1. Exploratory Colonial Period (1822 – 1909), and, 2. Expansionary Experimental Preand Post-Independence Period (1910 – 2010). During the first era which spanned early 19th to the end of the 19th century, cocoa seeds and plants were transported in barrels and shipments across the sea from the northern parts of Southern America and Central America to Africa. This was in response to the then imbibed and appreciated Aztec and Mayan culture of drinking '*chocolatl*', 'The Food of the Gods". The favourable similar tropical humid climatic conditions and abundant rainforest vegetation provided impetus for this transatlantic exploratory introduction which fortunately gave good results. The cultivation

**2. Cacao introduction history and genetic materials** 

economy, a status it still maintains today.

to the GDP.

of cocoa was therefore successfully established along the rainforest belt of West and Central African countries with significant economic revolution for both producing countries and chocolate lovers especially in the North. A timeline of earlier germplasm introduction efforts during the first era is presented in Table 2 below. The first materials introduced were of Amazonian origin, unlike the Criollo varieties introduced to Asiatic and Oceanic regions. The germplasm established on the island of Principe in 1822 was the main basis of the cocoa industry on the island. Original planting was said to have consisted of 30 plants which most likely were taken from a single fruit (pod). Progenies of these trees provided seeds for planting other areas of the island (Bartley 2005). Timeline of earliest cacao introduction into island and mainland West and Central Africa in the 19th and early 20th century



2. Sao Tome 'varieties' Sao Tome Introduction by Preuss,

1857 Seeds from Suriname Surinam Unsuccessful attempt by

Principe

Royal

1903 'Ocumare', 'Trinidad Trinidad Introduced plants were

BotanicGarden, England

Po/Sao Tome

(1901)

Royal Botanic Gardens, England

South and Central America (Trinidad)

Sao Tome Establishment of cocoa

Sao Tome

Tome

Trinidad Gosselin (1895), Preuss

2005)

1887.

Trinidad During this period,

in Fernando Po (now Bioko) with seeds from

13 plants shipment by a British missionary on Cameroon Mountains.

Curator of Victoria Botanic Garden. Some ofthese materials became known as 'Victoria-Kakao' variety of Sao

Introduction by Preuss. At the beginning of the 20th century, Cameroon had established the most diverse collection of cacao varieties (Bartley,

the Basel Missionaries.

Partial success by the Basel Missionaries.

introduction was to obtain varieties higher in quality than the widely grown 'Amelonado' from Fernando Po.

Tetteh Quarshie brought seeds from Fernando Po. Governor Griffiths also made introduction in

**Equatorial Guinea (Fernando Po)** 1854 Sao Tome 'Creoulo' and

1895? 332 plants including 'Forastero'

1900 Several varieties including 'Forastero', 'Criollo', 'Puerto-Cabello',

Colombia

1. 'Cundeamour', 'Pentagonum' from

Nicaragua

 2. 'Red Forastero', 'Criollo', 'White variety' of 'Caracas' type from

Jamaica

'Venezuela', 'Maracaibo', 'Guayaquil', 'La Guira', 'Soconusco', 'Suriname' and 'Nueva Grenada' from

1861 'Amelonado' Sao Tome &

'Amelonado' Fernando

**Cameroon**

**Ghana** 

1878? 1887

1900 - 1901

later, the 1880's Non-Amazonian 'other'varieties.

1876 1. Unknown materials likely from Trinidad.

Criollo', Nicaraguan Criollo', 'Red Criollo' and 'Yellow Criollo' established in Aburi Botanic Garden. These materials were largely self-incompatible. These are the likely ancestors of red fruited trees in Ghana. **Nigeria** *Regional differences in genetic composition of populations likely due to different routes of introduction A. The Niger Delta Protectorate covering Midwestern to Calabar Region*  1874 'Amelonado', Sao Tome 'Creuolo' Fernado Po Chief Squiss Ibaningo, a migrant worker introduced cocoa into Bonny, now in River State. This variety constituted virtually all the planting material cultivated at the middle of the 20th century. 1899 Unknown varieties. Forastero? Royal Botanic Garden, England Up to two shipments of cocoa plants 1900 'Pentagonum' variety Trinidad? Variety found at Old Calabar Station 1905 Non-Amelonado 'types' Trinidad? SaoTome? These were brought into Old calabar Station 1909 60 fruits of 'Forastero' type Trinidad Barrel shipment from Trinidad *B. The Lagos Colony* 1880? Sao Tome 'Comum' (Amelonado) Fernando Po First planting of cocoa in Lagos of some 1,500 plants. The plantation near Agege was owned by JPL Davies (Webster 1964) 1877-1888 Unknown. Fernando Po Trinidad Most likely non-Amelonado plants sent to Ceylon **Cote D'Ívoire**  1880 Amelonado Fernando Po First cocoa introduction **Sierra Leone**  Pre-1900 Sao Tome 'Comum' (Amelonado); Cameroon variety, Victoria-Kakao SaoTome, Fernando Po and Cameroon Several Sierra Leoneans were influential farmers in Fernando Po at the end of the 19th century. - 1902 - 60 plants of 'Ceylon Red', Trinidad


#### Table 2.

(Bartley 2005). By 1840, some quantities of cocoa was exported from the island. The cultivation of cocoa spread to the main island of Sao Tome in the 1850s. The variety which became known as Sao Tome "Creoulo", was self-compatible and homozygous and mostly related to the 'Comum'variety in Bahia, Brazil. This variety was taken both directly and indirectly through Fernando Po to other inland West and central African countries and became the basis of cocoa grown there. However, some other varieties were also introduced into Sao Tome from Ecuador, Trinidad and Venezuela in 1880. Consequently, the bulk of cocoa grown on farmers' plantation must have consisted of a mixture of these earlier varieties, but due to differential expression of self-incompatibility systems, the selfcompatible 'West African Amelonado' types must have dominated in the complex mixture of cacao of diverse origin at the beginning of the 20th century.

During the second era which began at about the end of the 19th century and beginning of the 20th century, economic considerations for higher income and premium due to greater yields and higher bean and chocolate quality were the main reasons for germplasm introduction. Previously selected individuals (clones) rather than 'types' showing potentials for high yields, resistance or tolerance to pests, diseases and abiotic stress such as drought were introduced and engaged in cultivar development processes on experimental stations. During this last decade, however, the "People, Planet and Profit'' concept of *Sustainability* has become a significant factor in cacao germplasm introduction. This has bearing with the concept of "Preventive Breeding" where clones showing resistance to regionally important diseases of cocoa growing regions could be introduced through international intermediate quarantine centers. This was to ensure that in the unlikely case of disease spread, for example, witches broom from South America to Africa, there is present in the African germplasm collections, sources of resistance to cope with the new disease in order not to paralyze the local cocoa economy as is the case during any outbreak.

Since the first successful introduction of *'Amelonado*' cacao, Lower Amazon Forastero type into West Africa in the late 19th century, there has been series of additional germplasm introductions as reviewed by (Bartley, 2005; Aikpokpodion, 2009). In Nigeria for instance, since formal selection and germplasm conservation programs around 1931 at the Nigerian Department of Agriculture in Moor Plantation, Ibadan there has been. Further germplasm introduction of Trinitario and Criollo selections from Trinidad and Ceylon (now Sri Lanka) (Jacobs et al., 1971). The British West African Colonial Administration established the West African Cocoa Research Institute (WACRI) in 1938 with headquarters in Tafo, Ghana and a mandate covering Gold Coast (Ghana), SierraLeone, Nigeria and Liberia. Several materials

(Bartley 2005). By 1840, some quantities of cocoa was exported from the island. The cultivation of cocoa spread to the main island of Sao Tome in the 1850s. The variety which became known as Sao Tome "Creoulo", was self-compatible and homozygous and mostly related to the 'Comum'variety in Bahia, Brazil. This variety was taken both directly and indirectly through Fernando Po to other inland West and central African countries and became the basis of cocoa grown there. However, some other varieties were also introduced into Sao Tome from Ecuador, Trinidad and Venezuela in 1880. Consequently, the bulk of cocoa grown on farmers' plantation must have consisted of a mixture of these earlier varieties, but due to differential expression of self-incompatibility systems, the selfcompatible 'West African Amelonado' types must have dominated in the complex mixture

During the second era which began at about the end of the 19th century and beginning of the 20th century, economic considerations for higher income and premium due to greater yields and higher bean and chocolate quality were the main reasons for germplasm introduction. Previously selected individuals (clones) rather than 'types' showing potentials for high yields, resistance or tolerance to pests, diseases and abiotic stress such as drought were introduced and engaged in cultivar development processes on experimental stations. During this last decade, however, the "People, Planet and Profit'' concept of *Sustainability* has become a significant factor in cacao germplasm introduction. This has bearing with the concept of "Preventive Breeding" where clones showing resistance to regionally important diseases of cocoa growing regions could be introduced through international intermediate quarantine centers. This was to ensure that in the unlikely case of disease spread, for example, witches broom from South America to Africa, there is present in the African germplasm collections, sources of resistance to cope with the new disease in order not to

Since the first successful introduction of *'Amelonado*' cacao, Lower Amazon Forastero type into West Africa in the late 19th century, there has been series of additional germplasm introductions as reviewed by (Bartley, 2005; Aikpokpodion, 2009). In Nigeria for instance, since formal selection and germplasm conservation programs around 1931 at the Nigerian Department of Agriculture in Moor Plantation, Ibadan there has been. Further germplasm introduction of Trinitario and Criollo selections from Trinidad and Ceylon (now Sri Lanka) (Jacobs et al., 1971). The British West African Colonial Administration established the West African Cocoa Research Institute (WACRI) in 1938 with headquarters in Tafo, Ghana and a mandate covering Gold Coast (Ghana), SierraLeone, Nigeria and Liberia. Several materials

Fernando Po

Sao Tome Many Liberians were

Tome.

contract workers in Sao

Nicaraguan Criollo', 'Forastero' and *T. pentagonum*

of cacao of diverse origin at the beginning of the 20th century.

paralyze the local cocoa economy as is the case during any outbreak.

 - 1000 seeds of unknown variety

1861? Sao Tome 'Comum' (Amelonado)

 Liberia

Table 2.

belonging to Upper Amazon Forastero and Trinitario populations were introduced from Trinidad by WACRI in 1944 (Toxopeus, 1964). Efforts to increase genetic variability in the base population in response to outbreaks of disease epidemic had provided impetus for germplasm introduction into Africa. For instance, the outbreak of cocoa swollen shoot disease in the 1930s in Ghana, Togo, and Nigeria almost destroyed the cocoa industry due to insufficient genetic variability in the base population. Consequently, new introductions were made in 1944 from Upper Amazon Forastero materials collected by F. J. Pound into the West African Cocoa Research Institute headquarters in Tafo, Ghana and Ibadan in Nigeria (Aikpokpodion et al., 2009). Due to the precocity of these materials, they were widely distributed for replanting of cut out plantations and by late 1950s, some 11 selected Upper Amazon types have been used to produce second and third generations of Amazon known as "F3 Amazon" or "Mixed Amazon" distributed to farmers (Knight and Rogers 1955). By 1961, some 60,000 ha in Ghana and an estimated 21 million seedlings had been distributed by the government of the Western Region to plant some 9,500 ha in Nigeria (Aikpokpodion 2009). Several hybrid varieties involving crosses with local Amelonado, Trinitario, and some Criollo materials were also developed from these materials in Ghana (Lockwood and Gyamfi 1979), Nigeria (Atanda and Jacobs 1974), and Cote d'Ivoire (Besse 1975; N'Goran et al. 1992).
