Losses of biodiversity and ecosystem functioning due to rainforest destruction and agricultural intensification are prime concerns for science and society alike. Potentially, ecosystems show nonlinear responses to land-use intensification that would open management options with limited ecological losses but satisfying economic gains. However, multidisciplinary studies to quantify ecological losses and socioeconomic tradeoffs under different management options are rare. Here, we evaluate opposing land use strategies in cacao agroforestry in Sulawesi, Indonesia, by using data on species richness of nine plant and animal taxa, six related ecosystem functions, and on socioeconomic drivers of agroforestry expansion. Expansion of cacao cultivation by 230% in the last two decades was triggered not only by economic market mechanisms, but also by rarely considered cultural factors. Transformation from near-primary forest to agroforestry had little effect on overall species richness, but reduced plant biomass and carbon storage by Ϸ75% and species richness of forest-using species by Ϸ60%. In contrast, increased land use intensity in cacao agroforestry, coupled with a reduction in shade tree cover from 80% to 40%, caused only minor quantitative changes in biodiversity and maintained high levels of ecosystem functioning while doubling farmers' net income. However, unshaded systems further increased income by Ϸ40%, implying that current economic incentives and cultural preferences for new intensification practices put shaded systems at risk. We conclude that low-shade agroforestry provides the best available compromise between economic forces and ecological needs. Certification schemes for shade-grown crops may provide a market-based mechanism to slow down current intensification trends.agricultural economics ͉ agroforestry management ͉ land use change ͉ plant-animal interactions ͉ ecosystem goods and services G lobal-scale conversion of tropical rainforests and agricultural intensification are major causes of biodiversity loss, and threaten ecosystem functioning, sustainable land use and local economies depending on natural resources (1-3). Developing strategies to reconcile human needs with the integrity of our environment is a major task for ecologists and socio-economists alike (4), but multitaxa studies are rare (5-6) and too little is known about the human dimension of land use changes (4, 7-11) and consequences for ecosystem functioning (1,2,(12)(13)(14). Furthermore, most ecological and economic studies on ecosystem services are carried out separately so that information cannot be brought together (15). Particularly, quantitative data on potential tradeoffs between biodiversity loss and agricultural intensification including natural habitat conversion is missing. Two competing solutions propose either wildlife-friendly farming on the cost of agricultural yields or land sparing by agricultural intensification to minimize the demand for natural habitat (16). The evaluation of such opposing land use options depends on t...

Although ecologists traditionally focus on natural ecosystems, there is growing awareness that mixed landscapes of managed and unmanaged systems provide a research environment for understanding basic ecological relationships on a large scale. Here, we show how tropical agroforestry systems can be used to develop ideas about the mechanisms that influence species diversity and subsequent biotic interactions at different spatial scales. Our focus is on tropical plantation crops, mainly coffee and cacao, and their pollinators, which are of basic ecological interest as partners in an important mutualistic interaction. We review how insect-mediated pollination services depend on local agroforest and natural habitats in surrounding landscapes. Further, we evaluate the functional significance of pollinator diversity and the explanatory value of species traits, and we provide an intercontinental comparison of pollinator assemblages. We found that optimal pollination success might be best understood as a consequence of niche complementarities among pollinators in landscapes harboring various species. We further show that small cavity-nesting bees and small generalist beetles were especially affected by isolation from forest and that larger-bodied insects in the same landscapes were not similarly affected. We suggest that mixed tropical landscapes with agroforestry systems have great potential for future research on the interactions between plants and pollinators.

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 α diversity (the number of species found per sampling unit) between taxa rather than on β 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 β diversity at local and landscape scales. We compared α and β 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 β 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 α diversity was not influenced by sampling completeness, whereas the indicator value for β diversity improved when using a similarity index that accounts for incomplete sampling. The indication potential of α diversity for β 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 α diversity and that patterns of β 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 β diversity shaping overall diversity patterns. Abstract. 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 a diversity (the number of species found per sampling unit) between taxa rather than on b 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 b diversity at local and landscape scales. We compared a and b 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 ...

The recent trend to place monetary values on ecosystem services has led to studies on the economic importance of pollinators for agricultural crops. Several recent studies indicate regional, long-term pollinator declines, and economic consequences have been derived from declining pollination efficiencies. However, use of pollinator services as economic incentives for conservation must consider environmental factors such as drought, pests, and diseases, which can also limit yields. Moreover, "flower excess" is a well-known reproductive strategy of plants as insurance against unpredictable, external factors that limit reproduction. With three case studies on the importance of pollination levels for amounts of harvested fruits of three tropical crops (passion fruit in Brazil, coffee in Ecuador, and cacao in Indonesia) we illustrate how reproductive strategies and environmental stress can obscure initial benefits from improved pollination. By interpreting these results with findings from evolutionary sciences, agronomy, and studies on wild-plant populations, we argue that studies on economic benefits from pollinators should include the total of ecosystem processes that (1) lead to successful pollination and (2) mobilize nutrients and improve plant quality to the extent that crop yields indeed benefit from enhanced pollinator services. Conservation incentives that use quantifications of nature's services to human welfare will benefit from approaches at the ecosystem level that take into account the broad spectrum of biological processes that limit or deliver the service.

No abstract

Throughout the tropics, agroforests are often the only remaining habitat with a considerable tree cover. Agroforestry systems can support high numbers of species and are therefore frequently heralded as the future for tropical biodiversity conservation. However, anthropogenic habitat modification can facilitate species invasions that may suppress native fauna. We compared the ant fauna of lower canopy trees in natural rainforest sites with that of cacao trees in agroforests in Central Sulawesi, Indonesia in order to assess the effects of agroforestry on occurrence of the Yellow Crazy Ant Anoplolepis gracilipes, a common invasive species in the area, and its effects on overall ant richness. The agroforests differed in the type of shadetree composition, tree density, canopy cover, and distance to the village. On average, 43% of the species in agroforests also occurred in the lower canopy of nearby primary forest and the number of forest ant species that occurred on cacao trees was not related to agroforestry characteristics. However, A. gracilipes was the most common non-forest ant species, and forest ant richness decreased significantly with the presence of this species. Our results indicate that agroforestry may have promoted the occurrence of A. gracilipes, possibly because tree management in agroforests negatively affects ant species that depend on trees for nesting and foraging, whereas A. gracilipes is a generalist when it comes to nesting sites and food preference. Thus, agroforestry management that includes the thinning of tree stands can facilitate ant invasions, thereby threatening the potential of cultivated land for the conservation of tropical ant diversity.

We studied species richness, composition and vertical distribution of epiphytic bryophytes in submontane rainforest of Central Sulawesi. Bryophytes were sampled on eight canopy trees and on eight trees in the forest understorey. Microclimate was measured at trunk bases and at crown bases. The total recorded number of 146 epiphytic bryophyte species is among the highest ever reported for tropical forests and underlines the importance of the Malesian region as a global biodiversity hotspot. Species composition differed significantly between understorey trees and canopy tree trunks on the one hand, and the forest canopy on the other. Fourty-five percent of the bryophyte species were restricted to canopy tree crowns, 12% to the understorey. Dendroid and fan-like species mainly occurred in the forest understorey whereas tufts were most species rich in the tree crowns. The findings reflect the different microclimatic regimes and substrates found in the understorey and in the forest canopy. The results indicate that assessments of the bryophyte diversity of tropical forests are inadequate when understorey trees and tree crowns are excluded.

The ongoing destruction of tropical rainforests has increased the interest in the potential value of tropical agroforests for the conservation of biodiversity. Traditional, shaded agroforests may support high levels of biodiversity, for some groups even approaching that of undisturbed tropical forests. However, it is unclear to what extent forest fauna is represented in this diversity and how management affects forest fauna in agroforests. We studied lower canopy ant and beetle fauna in cacao agroforests and forests in Central Sulawesi, Indonesia, a region dominated by cacao agroforestry. We compared ant and beetle species richness and composition in forests and cacao agroforests and studied the impact of two aspects of management intensification (the decrease in shade tree diversity and in shade canopy cover) on ant and beetle diversity. The agroforests had three types of shade that represented a decrease in tree diversity (high, intermediate and low diversity). Species richness of ants and beetles in the canopies of the cacao trees was similar to that found in lower canopy forest trees. However, the composition of ant and beetle communities differed greatly between the agroforest and forest sites. Forest beetles suffered profoundly from the conversion to agroforests: only 12.5% of the beetle species recorded in the forest sites were also found in the agroforests and those species made up only 5% of all beetles collected from cacao. In contrast, forest ants were well represented in agroforests, with 75% of all species encountered in the forest sites also occurring on cacao. The reduction of shade tree diversity had no negative effect on ants and beetles on cacao trees. Beetle abundances and non-forest ant species richness even increased with decreasing shade tree diversity. Thinning of the shade canopy was related to a decrease in richness of forest ant species on cacao trees but not of beetles. The contrasting responses of ants and beetles to shade tree management emphasize that conservation plans that focus on one taxonomic group may not work for others. Overall ant and beetle diversity can remain high in shaded agroforests but the conservation of forest ants and beetles in particular depends primarily on the protection of natural forests, which for forest ants can be complemented by the conservation of adjacent shaded cacao agroforests.