Abstract Arbuscular mycorrhizal fungi (AMF) play a central role in enhancing nutrient uptake by plants, increasing resilience to biotic and abiotic stresses, and supporting proper functioning of tropical agroecosystems, such as cacao plantations. However, little is known about diversity and capacity of native strains in rhizospheres, as well as soil factors shaping AMF communities. This study aimed to assess (i) AMF diversity and abundance in cocoa rhizospheres and the relationship with soil chemical properties and climatic variables at seven distinct areas in Côte d’Ivoire, i.e., Adzope, Agnibilekrou, Azaguie, Biankouma, Ble, Bonon and Soubre; (ii) in-situ capacity of strains from these rhizospheres. Soil samples were collected in 66 cocoa plots and AMF spores were extracted and identified based on morphometric criteria. Part of soils was used for trap cultures, while another part was used for determination of chemical properties. Our findings indicate that fungal communities associated with cacao are dominated by families Glomeraceae and Acaulosporaceae. Before establishment of trap cultures, cacao rhizospheres showed highest spore abundance at Ble (103 ± 47 spores/100 g soil) and lowest abundance at Biankouma (47.3 ± 21.6 spores/100 g soil). AMF strains from Agnibilekrou and Biankouma soils showed highest quantitative sporulation capacity, whereas strains from Soubré soils exhibited strongest infective capacity. 18.49% of variance in AMF community is explained by soil chemical variables, whereas 5.32% is explained by climatic variables, with Mn, Zn, Cr, and precipitation as main drivers. This study highlighted that a clear understanding of AMF communities in cocoa rhizospheres is essential to facilitate the design of efficient soil and plant conservation strategies, thus ensuring the sustainability and long-term productivity of cocoa agroecosystems in Côte d’Ivoire.

L’agroforesterie, souvent présentée comme solution à l’érosion de la biodiversité et aux chocs climatiques, échoue fréquemment lorsqu’elle repose sur des programmes massifs de plantation. Une autre approche, centrée sur la parcelle, valorise la complémentarité entre arbres rémanents, spontanés et plantés. Cette complémentarité renforce biodiversité, carbone, productivité et durabilité des cacaoyères.

Description of the subject. Tree diversity is essential for maintaining ecological richness and the survival of threatened species. Identifying the factors that favour it is therefore necessary to better guide agricultural policies. Objectives. The aim of this study is to analyse the determinants of tree diversity in the cocoa-growing areas of Côte d'Ivoire Method. The data used come from a survey of 150 cocoa farmers selected in the localities of Bonon, Biankouma and Soubré. These sites were chosen according to an East-West gradient corresponding to the evolution of the cocoa loop in Côte d'Ivoire. Floristic inventories were carried out in the cocoa farms of the selected farmers in order to determine floristic diversity. A Kruskal-Wallis test was performed to compare the farms and multiple linear regression was used to identify the determinants of this diversity. Results. The Biankouma zone has a higher diversity index, both regionally (5.12) and by plot (3.47), compared with the other two zones, the Soubré zone (4.69 regionally and 3.11 by plot) and the Bonon zone (4.22 regionally and 2.98 by plot). This result reflects a decline in floristic diversity as a function of the history of cocoa production. Regression analysis reveals that experience in cocoa production and the density of associated crops have a negative influence on farm diversity. Conversely, the number of associated tree species, the level of education and the distance separating the fields from nearby towns have a positive impact on this diversity. Conclusions. This study recommends promoting agroforestry to diversify cocoa farms. Agroforestry programmes should also focus on farms close to urban centres, to reconcile production goals with biodiversity conservation.

Cocoa cultivation in West Africa has been a major driver of deforestation, leading to increased greenhouse gas emissions and threatening cocoa yields. Agroforestry, which integrates trees from various origins-remnant, spontaneous, and planted-presents a sustainable solution to enhance carbon sequestration and improve farm resilience. However, the specific contributions of these tree origins and the socio-environmental factors shaping their effectiveness remain poorly understood. This study examines carbon dynamics across 150 cocoa fields in Côte d'Ivoire, analyzing a total of 11,568 trees across 15 sites. Using Bayesian modeling, we assess carbon stocks and gains from tree growth to explore how socio-environmental factors influence carbon balance in cocoa fields. Carbon stocks varied widely with remnant having the highest median carbon stocks (6.33 Mg/ha), followed by spontaneous (2.06 Mg/ha) and planted trees (1.53 Mg/ha). Carbon gains are similar for planted and spontaneous trees up to 7 years, but afterward, spontaneous trees grow faster (11.20 ± 0.87 kg/year) than planted ones (3.96 ± 0.5 kg/year). Carbon stocks rose with informed farmers and former forest use, but declined with higher cocoa density. Carbon gains at the tree level is primarily influenced by ownership and previous forest land use with positive effects, while cocoa density and annual temperature have negative effects. To maximize carbon sequestration and ensure the sustainable management of agroforestry systems, interventions should prioritize securing land tenure, enhancing farmer training in tree botany, and promoting the conservation of remnant and spontaneous trees.

This article examines the production costs of cocoa farming in Côte d’Ivoire, West Africa, taking into account the opportunity cost approach. To this end, surveys were conducted among 228 farmers in three regions, Bonon, Soubré and Biankouma, following an east–west gradient. The estimated costs of using family labor and land were based on the opportunity cost approach. The financial costs associated with production were also taken into account. Comparative analyses between different localities and cropping systems highlighted specific workload characteristics. Finally, a principal component analysis (PCA) was used to profile producers according to their income levels and profits. The findings showed that family labor was the main component of cocoa production costs. Prices paid to farmers did not always cover all production costs, with 38% of farmers producing at a loss, and this was contingent on the agro-ecological zone. Furthermore, the agroforestry system proved to be more economical in terms of labor than the full-sun system. These results underline the relevance of the opportunity cost approach in assessing production costs and setting cocoa selling prices. This should lead to a review of public price-setting mechanisms to ensure fair remuneration for family labor.

The impact of intra-plot multi shading on cocoa tree productivity is a widely exploited aspect of agroforestry practice. This study, conducted on 1,000 m2 experimental plots at three agroclimatic sites in Côte d’Ivoire (Azaguié, Divo, Soubré), mapped the spatial heterogeneity of shade provision using light intensity measurements and then calculated cocoa productivity over two years of data collection. A geostatic approach (ordinary kriging) was used to analyze the shade in agroforestry plots, which revealed significant heterogeneity. This heterogeneity was primarily attributed to the phenological diversity of the companion trees. The analysis revealed that low shading levels (approximately 30%) emerged as thresholds where cocoa productivity was highest. However, it was observed that elevated shading levels, exceeding 60%, resulted in a significant decline in productivity. Among the companion trees, Ceiba pentandra, Eleais guineensis , and Persea americana offer light to moderate shade. Conversely, certain trees that provide heavy shade, such as Mangifera indica , are identified as less beneficial because they significantly reduce cocoa productivity. ### Competing Interest Statement The authors have declared no competing interest.

This article examines production costs in cocoa farming in Ivory Coast, West Africa, taking opportunity costs into account. The data used come from surveys of 228 farmers in three areas of the country: Bonon, Soubré and Biankouma. The choice of these zones was based on an East-West gradient. The opportunity cost approach was used to estimate the cost of using family labor (FL) and land. Financial costs for the various production operations were also estimated. Comparative analyses between different localities and cropping systems were carried out to elucidate specificities in terms of workload. Finally, a Principal Component Analysis (PCA) was carried out to profile producers according to income level and associated profit. The results showed that FL is the main component of cocoa production costs. Prices paid to farmers do not always cover all production costs, and 38% of farmers produce at a loss. Production costs are contingent on agroecological zones. Moreover, agroforestry proved to be more economical in terms of labor than the full-sun system. These results underline the importance of taking opportunity costs into account when assessing production costs and setting cocoa prices. This could lead to a revision of pricing mechanisms to ensure fair remuneration for FL.

With about 46% of global production, C & ocirc;te d'Ivoire is the world's leading producer of cocoa beans. However, this production contributes to deforestation, exacerbating the effects of climate change. In response to this observation, this study aims to deepen knowledge on the contribution of agroforestry systems in cocoa production areas in C & ocirc;te d'Ivoire to atmospheric carbon storage. These main areas are the Centre-West, South-West, and West. In these areas, floristic richness was determined in 115 plots. Carbon stocks in living biomass, dead matter, and soil were evaluated. The dynamics of carbon stocks with age were also determined. The results revealed that the West area contains the most diversified cocoa agroforests, with 161 species compared to 71 and 119 in the Centre-West and South-West, respectively. Entandrophragma angolense, Nesogordonia papaverifera, and Sterculia oblonga, common to these areas, are on the IUCN Red List. Carbon stock varies by area, its history, the practices present, and especially the associated species. Thus, in the former cocoa production zone (Centre-West) and the current main production zone (South-West), Elaeis guineensis is the main carbon reservoir, with 25.576 tC.ha(-)1 in the Centre-West and 36.862 tC.ha(-)1 in the South-West. In the West, local trees form the main carbon reservoir with 11.701 tC.ha(-)1. The dynamics of total carbon stocks show heterogeneous changes in production areas according to the different stages of development of agroforestry systems. This is evidence of the complexity of carbon flow and the dynamics of cocoa systems, which are strongly influenced by the sociology of the producers.

Abstract West Africa, the largest cocoa‐producing region globally, has experienced significant deforestation in recent decades, leading countries to implement large‐scale agroforestry policies; however, most studies on farmers' adoption of agroforestry fail to consider the social (Who?), historical (When?), geographical (Where?), and ecological (What?) factors that influence their motivations to value trees. Drawing from a sample of 150 farmers responsible for the management of 12,096 trees, we quantified the motivations of farmers for 10 material and immaterial uses of trees and used a Bayesian modelling framework to explore the relative importance of the 4 W framework in explaining general motivations, specific to each use, as well as the varying levels of specialization in tree management strategies among farmers. The distribution of use values by category shows that the highest values are associated with (i) agronomic uses (such as shade for cocoa trees and soil fertilization), (ii) food for human consumption, (iii) social purposes, and (iv) medicinal uses. All four aspects of the 4 W framework significantly contribute to understanding farmers' deep motivations, while the influence of each 4 W determinant varies based on the specific material and immaterial uses being considered. The level of specialization or diversification in cocoa farmers' motivations is significantly influenced by their knowledge of tree species and cocoa tree density, with knowledgeable farmers exhibiting greater diversification, while higher cocoa tree density and the presence of remnant trees lead to more specialized strategies that hinder agroforestry adoption. From a political perspective, it is urgent that stakeholders involved in the promotion of agroforestry consider all dimensions of the farmer‐field system. The diversity of farmers' life histories (Who), of cultivated landscapes (Where), of field systems (What), and of time trajectories (When) present both constraints and opportunities with which farmers must contend to transition to much‐desired agroforestry systems. Read the free Plain Language Summary for this article on the Journal blog.

Agroforestry systems (AFS) are gaining major interest in cocoa producing countries. However, the analysis of competition effects in multispecific complex agricultural systems remains a key lock-in preventing both (i) a thorough understanding of resource partitioning, and (ii) the co-design of spatial patterns and management of cocoa AFS (cAFS) minimising competition. We sampled 48 cAFS 1000 m2 plots in Central Cameroon and used an individual based approach to analyse the effect neighbouring tree community composition had on coca tree pod production. First, we determined the distance up to which each neighbouring tree influenced the production of a given cocoa tree and checked for the significance and the magnitude of this effect. We then explored the trade-offs between cocoa tree production and the abundance of associated trees. Our results first underline a significant intra-specific competition within the cocoa tree populations studied, which would need to be thinned down to 800-1100 individuals per ha. We also found a general negative effect of associated trees kept or grown for their fruits (e.g. mango, traditional species, citrus trees). Those effects were stronger when the trees were positioned between 6 and 11 m away from the cocoa trees. These trees clearly impaired pod production, especially for small diameter cocoa trees. Palm trees, however, had a positive effect on cocoa pod production, with suitable densities modelled ranging from 0 to 240 individuals per ha. Finally, we found both positive and negative effects of associated forest trees on cocoa pod production, which varied with the distance separating them from the cocoa trees and the size of the cocoa trees. Overall, our analyses show that cocoa production in complex cAFS is influenced both by intra-and inter-specific interactions while it remains difficult to distinguish between potential intertwined effects and resource limitations.

Duru et al. (Agron Sustain Dev 35:1259-1281, 2015) highlighted a missing tool for studying and improving the performance of cropping systems in the transition to highly diversified agriculture. In response, this paper proposes a concept for designing, modeling, monitoring, and auditing desired ecosystem services, in intercropping and agroforestry systems. We have labelled this concept ESSU (Ecosystem Services functional Spatial Unit). It delimits the smallest spatial unit encompassing all the interacting species and other functional components (e.g., crops, trees, livestock, spontaneous vegetation, semi-natural habitats such as hedges, ditches, forest patches, and animals) that together provide a specified set of ecosystem services. The novel ESSU concept allows representation of an entire diversified agroecosystem by the repetition of the spatial unit that provides the same sets of targeted ecosystem services as the agroecosystem it represents. It can then be used for various activities, such as the (i) design of more efficient agroecological systems according to the targeted ecosystem services; (ii) rapid audit of farming practices for biodiversity/resilience across large tracts of farmland as part of achieving Sustainable Development Goal 2 targets of sustainable food systems; and (iii) modeling such diversified agroecosystems using a motif adapted to represent the targeted ecosystem services and the species spacing design. We demonstrate that the ESSU concept is highly flexible and applicable to a wide range of diversified agroecosystems, like arable intercropping, crop-tree intercropping, tree-tree agroforestry, and agro-pastoralism. We also show its relevance and suitability for representing temporal changes over 1 year, across several years, and over decades, indicating its generalizability and flexibility. We argue that ESSU could open new theoretical and practical research avenues for the study of diversified agroecosystems. Considered with all the knowledge available on practices, biodiversity, and ecosystem services, ESSU might provide a learning-support tool to fill the knowledge gap about relationships among practices, biodiversity, and associated ecosystem services.

Cocoa production has been one of the main drivers of forest loss in West Africa. In the resulting post-forest landscapes, agroforestry has often been recognised as a solution to reconcile the preservation of trees and agriculture. Thus, a large number of tree plantation programs have been carried out in cocoa fields. Despite these major investments, the success of these plantations as a tool for "reforestation" of landscapes and sustainable timber production has never really been evaluated in fields where remnant trees, spontaneous trees and (trans)planted trees coexist. To quantify the current and future timber resource, we inventoried all trees in 150 cocoa fields distributed along the bioclimatic and historical gradients of the cocoa production area of Côte d'Ivoire, the world's leading producer. Our results show that (i) 19.6% of all associated trees are timber species, (ii) in plots where farmers actually introduced trees by planting, only 13.1% of trees have been (trans)planted, (iii) 69.7% of the current timber volume comes from remnant trees and (iv) spontaneous trees constitute 77% of the future timber resource. Based on our results, we propose 23 species along with their cultivation methods for the renewal of timber resource in cocoa fields. Overall, our results show the failure of plantation programs in cocoa fields and suggest to bet on natural regeneration to sustainably provide timber wood. Consequently, private companies supplying trees to farmers should focus on species that are complementary to those already present in natural regeneration. At the landscape level, remnant trees and residual forests should be preserved to maintain propagule sources. Finally, investments in reforestation of cocoa fields should be redirected towards training small farmers in silvicultural management techniques such as assisted natural regeneration and tree pruning.

The demand for cocoa has increased over the past years following the growth of cocoa-based products linked to the rise in living standard in highly populated countries. Cacao industry is therefore currently facing the dilemma of producing more cacao while ensuring its sustainability. Cacao monocrops and agroforestry systems (AFS) are two contrasting ways to produce cocoa, yet their impact on yields, contribution to farmer livelihood, cocoa quality remains understudied. Therefore, we reviewed existing literature comparing (1) monocrop cacao farming systems with (2) simple or (3) complex AFS. We found 19 comparisons of the cocoa yields in monocrops and simple AFS and 20 comparisons of monocrop and complex AFS. Three main research findings derive from this work. First, in about one third of cases, cacao trees yield more (or equally) in AFS than in monocrops. However, when considering only simple AFS, cacao trees yield more or equal to cacao monocrop in 52% of the cases. Second, cocoa AFS yields an average of 14% less than cacao monocrop. Yet, on average simple AFS yielded 2% less than cacao monocrops. Finally, there are too little elements to draw conclusions about the nexus between cocoa quality and cacao tree cultivation system.

Combining mixed trees with cocoa trees in so-called cocoa agroforestry systems is emerging as a viable option for regenerating cocoa cropping in Africa. Pure cocoa crop stands currently prevail in these areas but are running out of steam. Technical solutions are therefore urgently needed to stabilize cocoa-growing areas, reduce pressure on forests and adapt to climate change. A balance can be struck in stands by combining cocoa trees with diverse trees chosen by farmers for their different uses, while maintaining a suitable and sustainable cocoa yield. Ecological services such as carbon storage and crop protection are co-benefits of this balance. Recent research in Cameroon highlights that cocoa agroforestry stands can be managed using a straightforward indicator—measurement of the basal area of cocoa trees and associated trees. This indicator could be adopted for sustainable cocoa production certification purposes, while the observed convergence between local know-how and scientific results could facilitate joint drawing up of technical recommendations.

L’agroforesterie, qui associe des arbres aux cacaoyers, apparaît comme une option crédible pour renouveler la cacaoculture en Afrique. La culture pure de cacaoyers, dominante, y est en effet à bout de souffle. Il est donc urgent d’identifier des solutions techniques pour stabiliser les zones cacaoyères, réduire la tension sur les forêts et s’adapter à l’évolution du climat. Un équilibre peut être trouvé entre les cacaoyers et un panachage d’arbres choisis par les agriculteurs pour leurs usages variés, tout en maintenant un rendement en cacao convenable sur le long terme. Cet équilibre assure aussi des services écologiques comme le stockage du carbone et la protection phytosanitaire. Des travaux récents menés au Cameroun montrent que le pilotage des cacaoyères agroforestières peut s’appuyer sur un indicateur simple, la mesure de la surface terrière des cacaoyers et celle des arbres associés. Cet indicateur pourrait être adopté pour la certification du cacao durable, tandis que la convergence constatée entre savoirs locaux et résultats scientifiques pourrait aider à co-construire les conseils techniques.

Past studies showed a gradual expansion of tree cover over savannah in the forest–savannah boundary zone of Cameroon (Gillet et al. 2001). While the encroachment of savannah by forest is more and more impeded by human activities, farmers have proven that afforestation at the border of the forest is achievable using cocoa and specific techniques to build up an associated tree canopy (Jagoret et al. 2012). Furthermore, mature cocoa agroforestry systems created on savannah (S-cAFS) and in forest (F-cAFS) seem to exhibit comparable multi-strata structure linked to a multi-purpose objective of farmers in terms of livelihoods and long-term sustainable management of cocoa. By combining measurements of cocoa production, litterfall and cycling, soil quality, carbon storage and tree species diversity along an age gradient (1 to 70 years), we showed that those variables in S- and F-cAFS generally tended to comparable levels after several decades. Results also emphasized the ability of S-cAFS to increase most of the ecosystem services (ES) although the time needed to reach levels found in F-cAFS varied strongly amongst variables (Nijmeijer et al. 2019a,b). We also compared the impact of five shade tree species (Canarium schweinfurthii, Dacryodes edulis, Milicia excelsa, Ceiba pentandra, Albizia adianthifolia) and unshaded conditions on soil functions and cocoa yield in relation to plant functional traits and leaf litterfall within 8 cocoa farms 20 to 60 years old (Sauvadet et al. 2020). While no difference in cocoa yields could be detected between the different tree species and unshaded conditions because of high variability of data, the effects on soil functions varied largely among species. Shade tree species with the most dissimilar litter traits to cocoa (cocoa showing the lowest leaf litter quality) showed the largest improvement of soil functions. Low litter recalcitrance was strongly associated with increases in soil fertility indicators such as N and P availability, while pH, soil C and N contents increased with litter Ca restitution. Improvements of soil functions were low under the two fruit trees (Canarium and Dacryodes), medium under the legume tree Albizia, and high under the two timber trees (Milicia and Ceiba). According to the tree species and nutrient, nutrient recycling through litterfall could barely to largely offset the nutrient removal by cocoa beans and husks. This study corroborates that the two tall timber trees (Milicia and Ceiba) are some of the most appreciated companion species for cocoa production by farmers, alleging desirable light shade, higher soil fertility and cocoa yield. In order to better assess the role of shade trees in these cocoa systems, future research will need to extend these approaches, especially to understand how plant diversity can help to adapt to climate change, including higher temperature and longer dry seasons.

La agroforestería, que asocia árboles con cacaoteros, parece ser una opción posible para renovar el cultivo del cacao en África. El cultivo puro de árboles de cacao, dominante, no es sostenible y el potencial productivo disminuye con el tiempo. Por lo tanto, es urgente encontrar soluciones técnicas que estabilicen las áreas de cultivo de cacao, reduzcan la presión sobre los bosques y se adapten al cambio climático. Se puede encontrar un equilibrio entre los árboles de cacao y una mezcla de árboles escogidos por los agricultores para sus diversos usos, manteniendo al mismo tiempo un rendimiento de cacao adecuado a largo plazo. Este equilibrio también asegura servicios ecológicos como el almacenamiento de carbono y la protección fitosanitaria. Un trabajo reciente llevado a cabo en Camerún muestra que el manejo de las plantaciones de cacao agroforestales puede basarse en un indicador simple, la medición del área basal de los árboles de cacao y la de los árboles asociados. Este indicador podría ser adoptado para la certificación de cacao sostenible, mientras que la convergencia observada entre el conocimiento local y los resultados científicos podría ayudar a construir conjuntamente el asesoramiento técnico.

Understanding the components of plantain productivity is important for contributing to the food security challenge in West and Central Africa. The purpose of this study was to assess how production system and cropping system characteristics affect plantain productivity. Interviews with farmers, dynamic measurements of cropped diversity, recording of management practices and characterization of the harvested bunches were used to characterize 25 plantain fields in the form of 54 factors and 5 dependent variables. The average bunch weight measured was 11.6 kg. The within-field variability of the bunch weights measured was 4.2 kg. The calculated mean plantain yield was 6.8 t/ha/year and varied between 1.1 and 18.8 t/ha/year depending on the fields studied. Harvested bunches amounted to only 34% of the field potential. Segmentation analyses (CART) of the fields studied and analyses of variance identified 12 factors strongly linked to bunch sizes and plantain field lifetime. The highest bunch weights were measured in fields belonging to farmers who participated in training and also applied herbicide and nitrogen fertilizers more frequently and at higher rates. These practices also increased within-field variability for bunch weights. Lastly, the management practices recorded showed an intensification of chemical inputs in traditional plantain-based cropping systems. These results, especially the high within-field and between-field variability for bunch weights, call for better quantify the impact of planting material quality and varietal mixture on plantain productivity into plantain-based cropping systems.

In Africa’s main cocoa producing countries, rehabilitation of old cocoa orchards is increasingly debated but rarely adopted. In Central Cameroon, rehabilitation practices are regularly set up in old cocoa-based agroforestry systems (cAFS). To better understand the impact of such practices we built a chronosequence of 40 cAFS. We carried out specific surveys with farmers on each plot in order to check for rehabilitation effects on cocoa stands and associated woody species (AWS). We found that cocoa trees represented on average 88.2% of woody individuals and increased with age (from 84.7 to 91.5%). The cocoa stand basal area (BA) share significantly increased with age and reached up to 40.2% in the oldest systems. Cocoa, fruit and forest trees mean BA increased with aging. They were on average of 6.5, 5.7 and 10.7 m2 ha−1 respectively. Six different architectural types, different from the theoretical architectural evolution of cocoa trees over time, were identified. Among them, type 4 characterized by several orthotropic suckers of differing ages, was found typical of farmers’ cutting back practices. Type 4 cocoa trees density increased over time and its BA represented on average 60% of cocoa stand BA in the oldest systems. Concomitantly, farmer’s management of AWS led to continuous evolution of the systems both in terms of density and species composition. Our results show that (i) permanent densification and cutting back practices (type 4) allow the rejuvenation of cocoa stands while increasing cocoa stands BA share; (ii) the continuous management of AWS by farmers is undertaken to favour cocoa trees share over time by limiting inter-specific competition and promoting complementarity between cocoa trees and AWS. We argue that such practices explain a fair part of the long-term sustainability observed in cAFS from Central Cameroon and represent a model from which new rehabilitation schemes could be inspired.