Cacao domestication II: progenitor germplasm of the Trinitario cacao cultivar

Cacao (Theobroma cacao L.) has been cultivated in Central America since pre-Columbian times. The type of cacao cultivated in this region was called Criollo; cacao populations from the Amazon basin were called Forastero. The type of Forastero most commonly cultivated until 1950 was named Amelonado. Historical data show Trinitario cacao to have originated in Trinidad, resulting from natural hybridisation between Criollo and Amelonado Forastero. Doubts persist on the source of the Amelonado Forastero involved in the origin of Trinitario; the Amelonado parent may have come from the Lower Amazon, the Orinoco or the Guyanas. Most of the cacao cultivated worldwide until 1950 consisted of Criollo, Trinitario and Amelonado. From the early 1950s, Forastero material collected in the Upper Amazon region during the 1930s and 1940s began to be employed in breeding programmes. To gain a better understanding of the origin and the genetic basis of the cacao cultivars exploited before the utilisation of germplasm collected in the Upper Amazon, a study was carried out using restriction fragment length polymorphism and microsatellite markers. Trinitario samples from 17 countries were analysed. With molecular markers, it was possible to clearly identify three main genotypes (represented by clones SP1, MAT1-6 and SIAL70) implicated in the origin of most Trinitario clones.     

Molecular Characterization of Cacao (<Emphasis Type="Italic">Theobroma cacao</Emphasis>) Germplasm from Jamaica Using Single Nucleotide Polymorphism (SNP) Markers

Cacao is an economically important commodity in Jamaica. Knowledge of the genetic diversity of Jamaican cacao germplasm is essential for their conservation and management. In spite of cacao’s economic importance in Jamaica, the crop is under studied, therefore limiting sound decisions toward improving productivity. Assessment of germplasm and on-farm genetic diversity is required to assist selecting superior genotypes to propagate and distribute across the island, as well as to use them as parental clones in breeding programs. Using 94 single nucleotide polymorphism (SNP) markers, 140 Jamaican cacao samples from two germplasm collections and a farmer’s estate along with 150 reference samples were analyzed. The principal coordinate analysis demonstrated that the majority of the Jamaican cacao selections were hybrids derived from five original germplasm groups, including Criollo, Amelonado and three Upper Amazon Forastero groups. Among the Upper Amazon groups, the Bayesian clustering analysis revealed that the Parinari (PA) ancestral lineage contributed the most (29.9%) to the Jamaican cacao germplasm. The germplasm collections showed greater diversity in terms of ancestral contributions compared to the farmer’s estate. However, the genetic differentiation between the three collecting sites was small (Fst?=?0.036), indicating that samples collected from the three sites were derived from a common pool of germplasm. The current study supports the historical records and clarified the ancestry of Jamaican cacao. Although the majority of the cacao genetic groups were observed in the Jamaican cacao collections, several diversity gaps were found in both germplasm collections and in the farmer’s estate, especially germplasm with disease resistance to cacao frosty pod rot that was recently found in Jamaica.     

Genetic identity and diversity of Nigerian cacao genebank collections verified by single nucleotide polymorphisms (SNPs): a guide to field genebank management and utilization

Nigeria is the sixth largest cacao producer in the world. Field performance and quality of cacao hybrid families is largely dependent on the genetic integrity of parental clones obtained in field genebank collections. However, information on the impact of mislabeling on seed garden output in Nigeria is lacking. Using 63 single nucleotide polymorphism (SNP) markers, we analyzed 1457 cacao trees sampled from seven major field genebank plots in Nigeria to assess the genetic integrity in Nigerian cacao germplasm. The procedure of multilocus matching with known reference clones revealed up to 78% mislabeling in recently introduced international germplasm. A high rate of mislabeling was also revealed in the West African local selections and breeding lines, using Bayesian assignment test. The problem of mislabeling has been attributed to errors from the sources of introduction, pre-planting labeling errors, and rootstocks overtaking budded scions due to poor field management. The analysis of genetic diversity revealed a good representation of the available cacao germplasm groups in Nigerian field genebanks, indicating that the genetic base of Nigeria cacao germplasm has been significantly widened through germplasm introductions. However, only a small proportion of the available germplasm in the genebank have been utilized for variety development. This study proved the utility of SNP markers for cleaning up the genebanks and reducing offtypes; thereby providing a strong basis for improving the accuracy and efficiency in cacao genebank management and breeding, as well as for mobilizing improved varieties to cacao farmers in Nigeria.     

Dissecting Genetic Structure in Farmer Selections of Theobroma Cacao in the Peruvian Amazon: Implications for on Farm Conservation and Rehabilitation

Knowledge of genetic diversity in farmers?? selections is essential for planning on-farm conservation and rehabilitation. Using 15 microsatellite loci, we analyzed parentage and population structure in 220 farmer selections of cacao from the Huallaga valley in Peruvian Amazon. A high level of allele richness and heterozygosity were detected in these selections. Coordination analysis showed that these farmer selections are mainly comprised of hybrids derived from Trinitario and Upper Amazon Forastero germplasm. Bayesian clustering analysis assigned 54 selections as Trinitario and 166 as Upper Amazon Forastero hybrids. Parentage analysis identified 15 international clones as probable parents for 96 farmer selections, which corresponded to a fraction of the known hybrid families disseminated in this region in the late 1980s. Combined analysis of demographic and molecular data revealed a significant spatial autocorrelation (r?=?0.235; P?=?0.006) at short geographical distances (<5.0 km). This patch-like distribution of spatial heterogeneity suggests a significant ??neighborhood effects?? in seeds distribution or variety adoption, where closely related hybrid progenies were adopted in the neighboring farms or villages. The outcomes of this study indicate that in spite of the introgressions of exotic germplasm in the past, Upper Amazon Forastero is still the dominant component in the Huallaga valley of Peru. The high level of on-farm diversity can offer needed variability for participatory selection of superior clones in this low input, small-scale production system, where adaptability to specific environment is more preferred than in a high input, large scale production system.     

Antimicrobial Effects of Ionizing Radiation on Artificially and Naturally Contaminated Cacao Beans

With an initial microbial level of ca. 10⁷ microorganisms per g of Ivory Coast cacao beans, 5 kGy of gamma radiation under an atmosphere of air reduced the microflora per g by 2.49 and 3.03 logs at temperatures of 35 and 50°C, respectively. Bahia cacao beans were artificially contaminated with dried spores of Aspergillus flavus and Penicillium citrinum, giving initial fungal levels of 1.9 × 10⁴ and 1.4 × 10³ spores per g of whole Bahia cacao beans, respectively. The average D₁₀ values for A. flavus and P. citrinum spores on Bahia cacao beans were 0.66 and 0.88 kGy, respectively.     

Establishment of a Core Collection of Traditional Cuban <Emphasis Type="Italic">Theobroma cacao</Emphasis> Plants for Conservation and Utilization Purposes

Presently, Theobroma cacao L. (cacao) in Cuba is mainly cultivated in the eastern region where plantations comprise a mixture of clonal varieties, hybrids, progeny of Trinidad Selected Hybrids, and traditional—also known as ancient—cacao. The ancient genetic resources, probably the plants most closely related to the original introductions, are endangered by their progressive replacement by modern clones. To promote the conservation and utilization of these genetic resources, a representative sample of 537 traditional Cuban cacao plants was used to develop a core collection. Core collections based on 15 simple sequence repeat (SSR) markers were generated using five different sampling algorithms: random sampling, simulated annealing, stepwise clustering with random sampling, the M strategy, and maximum genetic diversity. The five core collections were designed to capture 95 % of the SSR alleles in the complete collection. The genetic, morphological, and geographical diversity of each core collection was compared with that of the entire collection. The entire collection contained 139 alleles, including 104 rare ones, with the 95 % allelic coverage threshold achieved with 133 alleles. The core collection generated by the maximum genetic diversity algorithm had the lowest number of accessions (185), the highest mean genetic distance (0.486), the lowest morphological character redundancy, and the highest diversity as assessed by the mean Shannon-Weaver diversity index (0.757). This core collection can thus serve as the basis of future improvement programs based on local genetic resources.     

Combining-ability for cacao (Theobroma cacao L.) yield components under southern Bahia conditions

The objective of this study was to assess five cacao cultivars (selfs) and 20 hybrids with regard to their general-and specific-combining ability for yield components using method 1, model I, of the diallel analysis system. The selfings and the hybrids were obtained through controlled crossings, tested in the field in a random block design with four replications and plots containing 16 plants. The experiment was set up in the Centro de Pesquisas do Cacau, in Itabuna, Bahia, Brasil, in 1975. The characteristics studied were: the number of healthy and collected fruits per plant (NHFP and NCFP), the weight of humid seeds per plant and per fruit (WHSP and WHSF), and the percentage of diseased fruits per plant (PDFP), for 5 years (1986–1990). The F-test values, highly significant for general combining ability (GCA) and specific combining ability (SCA), demonstrated the existence of variability for both effects. However, the effects of SCA were greater than those of GCA, when compared in terms of the average squared effects. This condition held for the characteristics NHFP, NCFP and WHSP, which shows the relative importance of the non-additive genetic effects over the additive effects. The reciprocal effects did not show significance. Breeding methods which explore the additive portion of genetic variance should be employed for obtaining higher-yielding cacao and high seed weight. For this, the segregant populations should involve cultivars CEPEC 1, SIAL 169 and ICS 1. Combinations involving the cultivar ICS 1 presented the most favorable results for the characteristics WHSP and WHSF, where the hybrid SIAL 169 x ICS 1 and its reciprocal were outstanding.     

Genetic diversity and relationships of cacao (Theobroma cacao L.) in southern Mexico

 Neotropical tree crops are affected by a combination of biological and human factors that complicate the study of genetic diversity and crop evolution. Genetic diversity and relationships among southern Mexican populations and horticultural collections of Theobroma cacao (chocolate, cocoa, cacao) are examined in light of the agricultural practices of the Maya. Collections of cacao were obtained from the extremes of its geographic range including archeological sites in southern Mexico where cacao was first domesticated. Genetic diversity was assayed by 57 informative random amplified polymorphic DNA (RAPD) marker loci. A unique sample of the total diversity found in this study exists in the southern Mexican populations. These populations are significantly different from all other cacao with regards to their profile of RAPD bands, including the ‘criollo’ variety, their morphological and geographical group. A population of cacao found in a sinkhole (cenote) in northern Yucatan with genetic affinities to populations in Chiapas suggests the Maya maintained plants far away from their native habitat. This finding concurs with known agroforestry practices of the Maya. Modern efforts to increase germplasm of tropical tree crops such as cacao should carefully examine archeological sites where genetic diversity, either deliberately or by chance, was collected and maintained by ancient cultures. Received: 21 May 1997 / Accepted: 9 October 1997     

Shade cacao plantations (<Emphasis Type="Italic">Theobroma cacao</Emphasis>) and bat conservation in southern Bahia,Brazil

Nearly 40% of the remaining Atlantic Forest in southern Bahia state, Brazil, is a rustic agroforest of cacao (Theobroma cacao). These traditional shade plantations, locally known as ‘cabrucas’, are habitats for forest dwelling species. Here we investigated the potential role of this traditional crop as a support for bat assemblages in southern Bahia, establishing the extent to which the bat community structure found in this agricultural system is influenced by the proximity of forest tracts. We compared the bat community attributes (richness, diversity and dominance) and species-level response (capture frequency) between native forest tracts and shade cacao plantations located in two distinct landscapes, which varied in total amount of forest (patch sizes and total forested area) and in their proximity to forest tracts. The cabrucas contiguous to forest tracts showed a rich and abundant bat community, with samples showing capture rates, species richness, diversity and evenness significantly higher than those reported for forest tracts. This situation changes, however, when shade plantations are located at some distance from forest tracts ( > 1000 m). Bat communities in these isolated cabrucas are less diverse than those found in forests and nearby cabrucas, but in both cases, species usually associated with pristine habitats were found. These findings suggest that cabrucas per se are not forest surrogates, and their potential to harbor forest dwelling bat species is closely linked with the existence of nearby forest tracts that may act as a source for species populations. Therefore, the entire landscape should be considered for management, taking into account that maintenance of cabrucas together with the preservation and restoration of forest patches is probably direly needed if one wishes to conserve the bat diversity in southern Bahia for the long term.     

Diversity patterns of arbuscular mycorrhizal fungi associated with cacao in Venezuela

The arbuscular mycorrhizal fungal (AMF) communities associated with cacao in Venezuela were studied. The species of AMF spores present in sixteen cacao plantations and in one nursery were isolated and identified when possible. The spore densities, species richness, diversity, Shannon-Wiener diversity index and dominance concentration index for the AMF communities were calculated. Acaulospora scrobiculata was associated with cacao plants in all study sites. No Scutellospora spp. were found in the analyzed soils. The spore number found in cacao plantations was relatively lower as compared with other tropical crops (38 spores 100 g^–1 soil up to 1674). Soils that were cultivated with cacao for more than 40 years showed the lowest spore numbers. Species richness and diversity of AMF communities associated with cacao, were negatively correlated with available P in soils. The Shannon-Wiener diversity index was positively correlated with soil organic matter. These results indicate that the traditional cacao cultivation practices used in Venezuela, maintain mycorrhizal infection on cacao plants. The diversity of the AMF community is similar to that found in natural undisturbed ecosystems from Venezuela.     

Cacao Cultivation under Diverse Shade Tree Cover Allows High Carbon Storage and Sequestration without Yield Losses

One of the main drivers of tropical forest loss is their conversion to oil palm, soy or cacao plantations with low biodiversity and greatly reduced carbon storage. Southeast Asian cacao plantations are often established under shade tree cover, but are later converted to non-shaded monocultures to avoid resource competition. We compared three co-occurring cacao cultivation systems (3 replicate stands each) with different shade intensity (non-shaded monoculture, cacao with the legume Gliricidia sepium shade trees, and cacao with several shade tree species) in Sulawesi (Indonesia) with respect to above- and belowground biomass and productivity, and cacao bean yield. Total biomass C stocks (above- and belowground) increased fivefold from the monoculture to the multi-shade tree system (from 11 to 57 Mg ha^-1), total net primary production rose twofold (from 9 to 18 Mg C ha^-1 yr^-1). This increase was associated with a 6fold increase in aboveground biomass, but only a 3.5fold increase in root biomass, indicating a clear shift in C allocation to aboveground tree organs with increasing shade for both cacao and shade trees. Despite a canopy cover increase from 50 to 93%, cacao bean yield remained invariant across the systems (variation: 1.1–1.2 Mg C ha^-1 yr^-1). The monocultures had a twice as rapid leaf turnover suggesting that shading reduces the exposure of cacao to atmospheric drought, probably resulting in greater leaf longevity. Thus, contrary to general belief, cacao bean yield does not necessarily decrease under shading which seems to reduce physical stress. If planned properly, cacao plantations under a shade tree cover allow combining high yield with benefits for carbon sequestration and storage, production system stability under stress, and higher levels of animal and plant diversity.     

Tracing the native ancestors of the modern Theobroma cacao L. population in Ecuador

The native Theobroma cacao L. population from Ecuador, known as Nacional, is famous for its fine cocoa flavour. From the beginning of the twentieth century, however, it has been subjected to genetic erosion due principally to successive introductions of foreign germplasm whose hybrid descendants gradually replaced the native plantations, implying a decrease in cocoa quality. We attempted to trace this native cacao within a wide pool of modern Ecuadorian cacao population. Three hundred and twenty-two cacao accessions collected from different geographical areas along the pacific coast of Ecuador and maintained in two living collections were analysed using 40 simple-sequence repeat markers. Most of Ecuadorian cacao accessions displayed a high diversity and heterozygosity level. A factorial analysis of correspondence (FAC) showed a continuous variation among them, with a few ones, grouped at an extreme side of the FAC cloud, showing higher levels of homozygosity and lower introgression level by foreign cacaos. A paternity analysis revealed that these highly homozygous individuals are the most probable ancestors of the modern Nacional hybrid pool. These particular accessions studied could represent the native Nacional cacao present in Ecuador before the foreign introductions. Their identification will help to conserve valuable genetic material and to improve cocoa quality in new cacao varieties.     

Genetic diversity and structure of populations in Pilosocereus gounellei (F.A.C.Weber ex K.Schum.) (Cactaceae) in the Caatinga biome as revealed by heterologous microsatellite primers

Microsatellite transferability was used as a method to examine the genetic diversity and structure of populations in Pilosocereus gounellei seedling samples that have potential to implement effective restoration strategies for degraded and disturbed areas of the Caatinga biome. Genomic DNA was extracted from 85 seedlings obtained from fruit collected from plants growing in native areas in the Brazilian states of Piaui (PI), Rio Grande do Norte (RN), and Bahia (BA). Six microsatellite primers were polymorphic. AMOVA showed higher genetic variation within (72%) than among (28%) the samples from the three states. The high level of similarity between the seedlings from PI, BA, and RN indicated that samples collected at any of the three sites can be used to represent the genetic diversity of the species. Seeds of plants from the three States are recommended as samples for germplasm banks and/or the production of plantlets to i) plant in areas of strategic reserves for forage, ii) deploy new cultivation areas, iii) restore degraded areas in the semi-arid Northeast, and iv) maintain ecological reserve banks and fodder with genetically divergent plants.     

Conservation Value of Cacao Agroforestry Systems for Terrestrial Herbaceous Species in Central Sulawesi,Indonesia

Tropical secondary forest and agroforestry systems have been identified as important refuges for the local species diversity of birds and other animal groups, but little is known about the importance of these systems for terrestrial herbs. In particular, few studies report how the conversion from tropical forest to technified cacao plantation affects the species richness and the community structure of herbs. We conducted surveys in 43 cacao plantations along the border of the Lore Lindu National Park in Central Sulawesi, ranging from agroforests to technified cacao, categorizing the plantations as rustic cacao, planted shade cacao, and technified cacao. We recorded 91 herb species. Of the 74 species determined to species level, 21 were also found in natural forests, while 53 were recorded only in agricultural habitats. Araceae was the most forest‐dependent plant family while Asteraceae included the highest number of nonforest species. Overall, the presence of forest species was confined to moderately intensively managed rustic and planted shaded plantations. Distance from the forest, which has been identified as a crucial parameter for the diversity and composition of other taxa in cacao agroforests, only played a minimal role for herbs. Our study suggests that native forest herbs maybe more vulnerable to forest conversion than animal groups. The intensification of cacao plantation management increases the presence of weedy species to the detriment of native forest species.     

Microsatellite variation and population structure in the “Refractario” cacao of Ecuador

Utilization of germplasm for crop improvement is often hampered by absence of information regarding origin, genetic identity and genealogical relationships of germplasm groups or populations. Molecular marker technology offers an efficient tool to verify or reconstruct passport data. Using a high-throughput genotyping system with 15 microsatellite loci, we fingerprinted 482 accessions in 48 putative half-sib families of Refractario cacao (a group of germplasm collected from nine farms in Ecuador). Based on the multilocus profiles, a Bayesian method for individual assignment was applied to verify membership in each half-sib family. Multivariate statistical analysis showed that the Refractario genetic profile was different from other groups tested, except for the “Nacional” cacao from the coastal valley of Ecuador. Hierarchical partitioning of genetic variance in the Refractario cacao showed that 76% of the variation was contributed by intra-family difference, whereas the inter-family and inter-farm difference accounted for 15 and 9% of total variance, respectively. All three sources of variation were highly significant (P < 0.01). Cluster and Principal Coordinates Analyses revealed a population sub-structure in Refractario, which was also highly heterozygous, suggesting hybridization derived from Nacional cacao and multiple other parental varieties, which all shared a similar genetic background. The improved understanding of identities and structure in Refractario cacao will contribute to more efficient conservation and use of this germplasm group in cacao breeding.     

Conservation in tropical landscape mosaics: the case of the cacao landscape of southern Bahia, Brazil

A recent debate has contrasted two conservation strategies in agricultural landscapes; either “land sparing” farm development combining intensive production practices with forest set-asides, or “wildlife-friendly” farming with greater on-farm habitat value but lower yields. We argue that in established mosaic landscapes including old cacao production regions where natural forest has already been reduced to relatively small fragments, a combination of both strategies is needed to conserve biodiversity. After reviewing the evidence for the insufficiency of either strategy alone if applied to such landscapes, the paper focuses on the cacao production landscape of southern Bahia, Brazil, once the world’s second largest cacao producer. Here, small remaining areas of Atlantic Forest are embedded in a matrix dominated by traditional cacao agroforests, resulting in a landscape mosaic that has proven favorable to the conservation of the region’s high biodiversity. We show that current land use dynamics and public policies pose threats but also offer opportunities to conservation and describe a three-pronged landscape conservation strategy, consisting of (i) expansion of the protected areas system, (ii) promotion of productive yet biodiversity-friendly cacao farming practices, and (iii) assistance to land users to implement legally mandated on-farm reserves and voluntary private reserves. We discuss recent experiences concerning the implementation of this strategy, discuss likely future scenarios, and reflect on the applicability of the Bahian experience to biodiversity rich cacao production regions elsewhere in the tropics.     

Genetic variability in wild genotypes of Passiflora cincinnata based on RAPD markers

The genetic diversity and characteristics of commercial interest of Passiflora species make it useful to characterize wild germplasm, because of their potential use for fruit, ornamental and medicinal purposes. We evaluated genetic diversity, using RAPD markers, of 32 genotypes of Passiflora cincinnata collected from the wild in the region of Vitória da Conquista, Bahia, Brazil. Thirteen primers generated 95 polymorphic markers and only one monomorphic marker. The mean genetic distance between the genotypes estimated by the complement of the Dice index was 0.51 (ranging from 0.20-0.85), and genotype grouping based on the UPGMA algorithm showed wide variability among the genotypes. This type of information contributes to identification and conservation of the biodiversity of this species and for the identification of pairs of divergent individuals for maximum exploitation of existing variability.     

Ferns,frogs, lizards,birds and bats in forest fragments and shade cacao plantations in two contrasting landscapes in the Atlantic forest,Brazil

The traditional shade cacao plantations (cabrucas) of southern Bahia, Brazil, are biologically rich habitats, encompassing many forest-dwelling species. However, a critical question for the conservation management of this specific region, and the highly fragmented Atlantic forest in general, is to what extent the conservation value of cabrucas relies on the presence of primary forest habitat in the landscape. We investigated the relative importance of cabrucas and forests for the conservation of five diverse biological groups (ferns, frogs, lizards, birds and bats) in two contrasting landscapes in southern Bahia, one dominated by forest with some interspersed cabrucas, and one dominated by cabrucas with interspersed forest fragments. The community structure (richness, abundance and diversity) of all biological groups differed between cabrucas and forests, although these differences varied among groups. A high number of forest species was found in the cabrucas. However, there were pronounced differences between the two landscapes with regard to the ability of cabrucas to maintain species richness. Irrespective of the biological group considered, cabrucas located in the landscape with few and small forest fragments supported impoverished assemblages compared to cabrucas located in the landscape with high forest cover. This suggests that a greater extent of native forest in the landscape positively influences the species richness of cabrucas. In the landscape with few small forest fragments interspersed into extensive areas of shade cacao plantations, the beta diversity of birds was higher than in the more forested landscape, suggesting that forest specialist species that rarely ventured into cabrucas were randomly lost from the fragments. These results stress both the importance and the vulnerability of the small forest patches remaining in landscapes dominated by shade plantations. They also point to the need to preserve sufficient areas of primary habitat even in landscapes where land use practices are generally favorable to the conservation of biodiversity.     

Conservation of Vascular Epiphyte Diversity in Shade Cacao Plantations in the Chocó Region of Ecuador

To assess the contributions of rustic shade cacao plantations to vascular epiphyte conservation, we compared epiphyte species richness, abundance, composition, and vertical distributions on shade trees and in the understories of six plantations and adjacent natural forests. On three phorophytes and three 10 × 10 m understory plots in each of the agroforestry plantations and natural forests, 54 and 77 species were observed, respectively. Individual-based rarefaction curves revealed that epiphyte species richness was significantly higher on forest phorophytes than on cacao farm shade trees; detailed analyses showed that the differences were confined to the inner and outer crown zones of the phorophytes. No differences in epiphyte species richness were found in understories. Araceae, Piperaceae, and Pteridophyta were less species-rich in plantations than in forests, while there were no differences in Orchidaceae and Bromeliaceae. Regression analysis revealed that epiphyte species richness on trunks varied with canopy cover, while abundance was more closely related to soil pH, canopy cover, and phorophyte height. For crown epiphytes, phorophyte diameter at breast height (dbh) explained much of the variation in species richness and abundance. There were also pronounced downward shifts in the vertical distributions of epiphyte species in agroforests relative to natural forests. The results confirm that epiphyte diversity, composition, and vertical distributions are useful indicators of human disturbance and showed that while the studied plantations serve to preserve portions of epiphyte diversity in the landscape, their presence does not fully compensate for the loss of forests.     

Recovery of species richness and conservation of native Atlantic forest trees in the cacao plantations of southern Bahia in Brazil

The Atlantic forests of southern Bahia in Brazil present great species richness and a high degree of endemism. A large part of these native forests were transformed into cacao plantations in an agroforestry system known locally as cabrucas, where native trees were culled and cacao was planted under the shade of remaining trees. The present study analyzed the influence of time of implantation (age) and time of abandonment of management practices on tree species diversity of cabruca plantations to evaluate the capacity for conservation and recovery of species richness of native Atlantic Forest trees in cabrucas. Phytosociological surveys were conducted in five cabrucas with different conditions of age and state of abandonment. All trees, including hemiepiphytes and excluding the cacao plants, with a minimum stem diameter of 10 cm at breast height, were surveyed within a 3-ha sampling area in each plantation. A total of 2514 individual trees belonging to 293 species and 52 families were recorded in the five cabrucas. The Shannon diversity index varied from 3.31 to 4.22 among the cabrucas and was positively correlated with the time of abandonment (r = 0.97). The new cabrucas showed the highest values of estimated total richness (Chao) and the highest proportion of late successional species than the old ones. All areas preserved a very high proportion of native forest species while the three old cabrucas showed a higher proportion of exotic species than the two new ones. Thus the exotic species seem to replace more of the native species in the long run because of management practices and local preferences. The cabrucas presented also a high capacity for the regeneration of tree species richness after abandonment. Simple alterations in management practices could improve the recruitment of late successional species in these areas. Economic incentives may be necessary for the farmers to adopt management practices to retain native species which bring no economic returns.