Damayanti Buchori

Combining high biodiversity with high yields in tropical agroforests

Local and landscape-scale agricultural intensification is a major driver of global biodiversity loss. Controversially discussed solutions include wildlife-friendly farming or combining high-intensity farming with land-sparing for nature. Here, we integrate biodiversity and crop productivity data for smallholder cacao in Indonesia to exemplify for tropical agroforests that there is little relationship between yield and biodiversity under current management, opening substantial opportunities for wildlife-friendly management. Species richness of trees, fungi, invertebrates, and vertebrates did not decrease with yield. Moderate shade, adequate labor, and input level can be combined with a complex habitat structure to provide high biodiversity as well as high yields. Although livelihood impacts are held up as a major obstacle for wildlife-friendly farming in the tropics, our results suggest that in some situations, agroforests can be designed to optimize both biodiversity and crop production benefits without adding pressure to convert natural habitat to farmland.

Alpha and beta diversity of plants and animals along a tropical land-use gradient

Assessing the overall biological diversity of tropical rain forests is a seemingly insurmountable task for ecologists. Therefore, researchers frequently sample selected taxa that they believe reflect general biodiversity patterns. Usually, these studies focus on the congruence of alpha diversity (the number of species found per sampling unit) between taxa rather than on beta diversity (turnover of species assemblages between sampling units). Such approaches ignore the potential role of habitat heterogeneity that, depending on the taxonomic group considered, can greatly enhance beta diversity at local and landscape scales. We compared alpha and beta diversity of four plant groups (trees, lianas, terrestrial herbs, epiphytic liverworts) and eight animal groups (birds, butterflies, lower canopy ants, lower canopy beetles, dung beetles, bees, wasps, and the parasitoids of the latter two) at 15 sites in Sulawesi, Indonesia, that represented natural rain forest and three types of cacao agroforests differing in management intensity. In total, we recorded 863 species. Patterns of species richness per study site varied strongly between taxonomic groups. Only 13-17% of the variance in species richness of one taxonomic group could be predicted from the species richness of another, and on average 12-18% of the variance of beta diversity of a given group was predicted by that in other groups, although some taxon pairs had higher values (up to 76% for wasps and their parasitoids). The degree of congruence of patterns of alpha diversity was not influenced by sampling completeness, whereas the indicator value for beta diversity improved when using a similarity index that accounts for incomplete sampling. The indication potential of alpha diversity for beta diversity and vice versa was limited within taxa (7-20%) and virtually nil between them (0-4%). We conclude that different taxa can have largely independent patterns of alpha diversity and that patterns of beta diversity can be more congruent. Thus, conservation plans on a landscape scale need to put more emphasis on the high heterogeneity of agroforests and the overarching role of beta diversity shaping overall diversity patterns.

Ecological and socio-economic functions across tropical land use systems after rainforest conversion

Tropical lowland rainforests are increasingly threatened by the expansion of agriculture and the extraction of natural resources. In Jambi Province, Indonesia, the interdisciplinary EFForTS project focuses on the ecological and socio-economic dimensions of rainforest conversion to jungle rubber agroforests and monoculture plantations of rubber and oil palm. Our data confirm that rainforest transformation and land use intensification lead to substantial losses in biodiversity and related ecosystem functions, such as decreased above- and below-ground carbon stocks. Owing to rapid step-wise transformation from forests to agroforests to monoculture plantations and renewal of each plantation type every few decades, the converted land use systems are continuously dynamic, thus hampering the adaptation of animal and plant communities. On the other hand, agricultural rainforest transformation systems provide increased income and access to education, especially for migrant smallholders. Jungle rubber and rubber monocultures are associated with higher financial land productivity but lower financial labour productivity compared to oil palm, which influences crop choice: smallholders that are labour-scarce would prefer oil palm while land-scarce smallholders would prefer rubber. Collecting long-term data in an interdisciplinary context enables us to provide decision-makers and stakeholders with scientific insights to facilitate the reconciliation between economic interests and ecological sustainability in tropical agricultural landscapes.

Can joint carbon and biodiversity management in tropical agroforestry landscapes be optimized

Managing ecosystems for carbon storage may also benefit biodiversity conservation, but such a potential ‘win-win’ scenario has not yet been assessed for tropical agroforestry landscapes. We measured above- and below-ground carbon stocks as well as the species richness of four groups of plants and eight of animals on 14 representative plots in Sulawesi, Indonesia, ranging from natural rainforest to cacao agroforests that have replaced former natural forest. The conversion of natural forests with carbon stocks of 227–362 Mg C ha−1 to agroforests with 82–211 Mg C ha−1 showed no relationships to overall biodiversity but led to a significant loss of forest-related species richness. We conclude that the conservation of the forest-related biodiversity, and to a lesser degree of carbon stocks, mainly depends on the preservation of natural forest habitats. In the three most carbon-rich agroforestry systems, carbon stocks were about 60% of those of natural forest, suggesting that 1.6 ha of optimally managed agroforest can contribute to the conservation of carbon stocks as much as 1 ha of natural forest. However, agroforestry systems had comparatively low biodiversity, and we found no evidence for a tight link between carbon storage and biodiversity. Yet, potential win-win agroforestry management solutions include combining high shade-tree quality which favours biodiversity with cacao-yield adapted shade levels.

Land-use choices follow profitability at the expense of ecological functions in Indonesian smallholder landscapes

Smallholder-dominated agricultural mosaic landscapes are highlighted as model production systems that deliver both economic and ecological goods in tropical agricultural landscapes, but trade-offs underlying current land-use dynamics are poorly known. Here, using the most comprehensive quantification of land-use change and associated bundles of ecosystem functions, services and economic benefits to date, we show that Indonesian smallholders predominantly choose farm portfolios with high economic productivity but low ecological value. The more profitable oil palm and rubber monocultures replace forests and agroforests critical for maintaining above- and below-ground ecological functions and the diversity of most taxa. Between the monocultures, the higher economic performance of oil palm over rubber comes with the reliance on fertilizer inputs and with increased nutrient leaching losses. Strategies to achieve an ecological-economic balance and a sustainable management of tropical smallholder landscapes must be prioritized to avoid further environmental degradation.

Biodiversity patterns and trophic interactions in human-dominated tropical landscapes in Sulawesi (Indonesia): plants, arthropods and vertebrates

The need to capture primary production in order to sustain and improve economic livelihoods has lead to increasing conversion of natural habitat and intensification of agricultural practices in many parts of the world including most tropical regions. Understanding how these processes affect ecosystems and their functioning, in particular in the high-diversity ecosystems of the tropics, has become a key issue in ecological research. In this chapter, our focus is on the agriculture-forest landscapes of Central Sulawesi, Indonesia, an island widely known for its endemic yet still poorly known flora and fauna. The rise of the region to one of the largest cacao producing areas in the world is at the core of recent land-use change and intensification processes. Covering plants (trees, rattan palms, herbs, bryophytes) and several invertebrate (ants, dung beetles, cacao insect herbivores, fruit-feeding butterflies, parasitic Hymenoptera, spiders) and vertebrate groups (amphibians, birds, murids, reptiles), we give an in-depth overview of the determinants of biodiversity in cacao landscapes, including both management and landscape-scale variables into our analyses. Results show that shaded agroforests host a rich community of species. By adopting a large-scale study design we showed that proximity of natural forest is a key predictor for species richness of plants, invertebrates and vertebrates alike. Endemics and forest specialists benefit most from indigenous shade tree cover and proximity to natural forest. Importantly, several functionally important groups such as insectivorous and seed-dispersing birds benefit from tall shade trees, shade tree diversity and proximity to forest edge, while parasitoid diversity is greatest close to natural forests. Available data on the effects of landuse change in cacao landscape of Central Sulawesi is increasing. Change in landscape configuration and management practices are being clearly reflected in the composition of species communities, with likely impacts on ecosystem services such as pest control and pollination. More knowledge is needed especially in terms of species interactions and ecosystem functioning, but also on how existing knowledge can contribute to effective conservation in human-dominated landscapes outside protected areas.

Long-term change of ant community structure in cacao agroforestry landscapes in Indonesia

Land‐use change and agricultural intensification can strongly affect biodiversity in agricultural landscapes. Although many studies investigate management impacts, data on the long‐term change of species communities in agroecosystems are scarce. We analysed the long‐term change in diversity and composition of ant communities in cacao agroforestry systems in Central Sulawesi, Indonesia and attempted to disentangle the driving factors of this change. Ant communities were resampled in 2009 from sites for which previous surveys had been conducted either in 2001 (the rainforest‐poor Palolo region) or 2003 (the rainforest‐rich Kulawi region) using insecticide fogging. Ant community composition changed significantly over time in Palolo and Kulawi. The change in ant species richness differed between regions. Species richness increased in Kulawi, probably due to the increasing availability of nest sites and microhabitats as trees grow larger and older. In the Palolo region, species richness decreased, suggesting that the high local intensification and landscape‐wide changes may have counteracted the effects of tree age. Changes in ant communities over time were significant, but were more difficult to explain than expected, despite clear difference in management changes within and between regions. The findings suggest that the landscape‐scale differences between the two study regions play a more important role for species diversity and its composition than changes in local management. This highlights the importance of long‐term studies across contrasting landscapes for better understanding the consequences land‐use intensification has on tropical biodiversity.

Insect diversity responses to forest conversion and agroforestry management

The ongoing loss of pristine tropical rainforests increases the potential importance of agroforestry systems for the conservation of tropical arthropod diversity. Shaded agroforestry systems can still support high levels of biodiversity, even resembling those supported by undisturbed forests, but intensively managed open agroforestry systems may cause severe losses in insect diversity. In this study we evaluate the conservation value of agroforestry systems for species richness and diversity (Simpson’s index) of four insect groups at natural forest sites and three different types of cacao-dominated agroforestry systems in Central Sulawesi, Indonesia. The agroforestry systems were characterised by low, intermediate and high diversity of shade trees. Each habitat type was studied with 4 replicates, i.e. 16 study sites altogether. We compared responses of solitary bees and wasps, dung beetles and lower canopy dwelling beetles and ants. These taxa represent diverse and functionally important insect groups: solitary bees and wasps act as crop pollinators or pest predators, dung beetles as decomposers of mammalian excrements and canopy dwelling beetles and ants include abundant herbivores and predators. High percentages of forest species did not occur in agroforestry systems, but diversity and species richness in agroforests remained as high as or even higher than in the forest lower canopy. Diversity, species richness and abundance of the functionally important dung beetles and canopy ants showed strong resilience against both forest conversion and changes in agroforestry management. Diversity, species richness and abundance of solitary bees and wasps as well as canopy beetles even seemed to profit from the effects of opening the upper canopy that was related to forest conversion and changes in shade tree compositions. On the latter two groups the effects of opening the upper canopy were stronger than effects of reduced shade tree richness. Based on our results, we recommend the inclusion of agroforestry systems with a diversity of shade trees in tropical conservation plans in addition to pristine forest reserves. Furthermore, regional differences in local agroforestry management contributed to between 31% (for dung beetles) and 58% (for canopy beetles) of the total species richness, which stresses the importance of conservation policies aimed at a diversity of habitat types on a broader landscape scale.

Experimental Biodiversity Enrichment in Oil-Palm-Dominated Landscapes in Indonesia

Tropical biodiversity is threatened by the expansion of oil-palm plantations. Reduced-impact farming systems such as agroforests, have been proposed to increase biodiversity and ecosystem functioning. In regions where oil-palm plantations already dominate the landscape, this increase can only be achieved through systematic ecological restoration. However, our knowledge about the underlying ecological and socio-economic processes, constraints, and trade-offs of ecological restoration in oil-palm landscapes is very limited. To bridge this gap, we established a long-term biodiversity enrichment experiment. We established experimental tree islands in a conventional oil-palm plantation and systematically varied plot size, tree diversity, and tree species composition. Here, we describe the rationale and the design of the experiment, the ecosystem variables (soil, topography, canopy openness) and biotic characteristics (associated vegetation, invertebrates, birds) of the experimental site prior to the establishment of the experiment, and initial experimental effects on the fauna. Already one year after establishment of the experiment, tree plantings had an overall positive effect on the bird and invertebrate communities at the plantation scale. The diversity and abundance of invertebrates was positively affected by the size of the tree islands. Based on these results, we expect a further increase of biodiversity and associated ecological functions in the future. The long-term interdisciplinary monitoring of ecosystem variables, flora, fauna, and socio-economic aspects will allow us to evaluate the suitability of tree islands as a restoration measure. Thereof, guidelines for ecologically improved and socio-economically viable restoration and management concepts could be developed.

Spatiotemporal density patterns of the pest predator Rhynchium haemorrhoidale (F.) along a land-use gradient in cacao agroforestry systems

Tropical insect species show year-round breeding activity due to favourable climatic conditions. However, most species also display seasonal reproductive peaks, but little is known about underlying causes of temporal density changes. We investigated population dynamics of the pest predator Rhynchium haemorrhoidale (F.) (Hymenoptera: Eumenidae) and its natural enemies in relation to season, climate and varying shade tree composition in cacao agroforestry systems in Central Sulawesi (Indonesia). Nesting of R. haemorrhoidale showed clear seasonality with highest densities in the wet season and lowest in the dry season, which was not related to changes in temperature. Wasp densities increased with land-use intensity, presumably because less-shaded areas offer more favourable climatic conditions and higher densities of the major prey, the cacao pest Agathodes caliginosalis (Lepidoptera: Pyralidae). Pupal body size was higher in June than in November indicating increasing intraspecific competition due to high wasp densities and food shortage at the beginning of the wet season. Body size between habitat types was similar, thus season appeared to be more important than habitat in terms of food supply. High wasp densities in the wet season were associated with high diversity of the altogether seven parasitoid species. In conclusion, we found a pronounced seasonality of the wasp R. haemorrhoidale and a preference for little shaded agroforestry, which may be due to enhanced temperature and density of its prey. Our data indicate a high potential for biological control of a major cacao pest by management of nesting sites of R. haemorrhoidale.