Owing to complex direct and indirect effects, impacts of higher trophic levels on plants is poorly understood. In tropical agroecosystems, ants interact with crop mutualists and antagonists, but little is known about how this integrates into the final ecosystem service, crop yield. We combined ant exclusion and introduction of invasive and native-dominant species in cacao agroecosystems to test whether (i) ant exclusion reduces yield, (ii) dominant species maximize certain intermediate ecosystem services (e.g. control of specific pests) rather than yield, which depends on several, cascading intermediate services and (iii) even, species-rich ant communities result in highest yields. Ants provided services, including reduced leaf herbivory and fruit pest damage and indirect pollination facilitation, but also disservices, such as increased mealybug density, phytopathogen dissemination and indirect pest damage enhancement. Yields were highest with unmanipulated, species-rich, even communities, whereas ant exclusion decreased yield by 27%. Introduction of an invasivedominant ant decreased species density and evenness and resulted in 34% lower yields, whereas introduction of a non-invasive-dominant species resulted in similar species density and yields as in the unmanipulated control. Species traits and ant community structure affect services and disservices for agriculture in surprisingly complex ways, with species-rich and even communities promoting highest yield.

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 ...

Summary1. Avian ecosystem services such as the suppression of pests are considered to be of high ecological and economic importance in a range of ecosystems, especially in tropical agroforestry. However, how bird predation success is related to the diversity and composition of the bird community, as well as local and landscape factors, is poorly understood. 2. We quantified arthropod predation in relation to the identity and diversity of insectivorous birds using experimental exposure of artificial, caterpillar-like prey in 15 smallholder cacao agroforestry systems differing in local shade-tree management and distance to primary forest. The bird community was assessed using both mist-netting (targeting active understorey insectivores) and point counts (higher completeness of species inventories). 3. Bird predation was not related to local shade-tree management or overall bird species diversity, but to the activity of insectivorous bird species and the proximity to primary forest. Insectivore activity was best predicted by mist-netting-based data, not by point counts. We identified the abundant Indonesian endemic lemon-bellied white-eye Zosterops chloris as the main driver of predation on artificial prey. 4. Synthesis and applications. The suppression of arthropods is a major ecosystem service provided by insectivorous birds in agricultural systems world-wide, potentially reducing herbivore damage on plants and increasing yields. Our results show that avian predation success can be driven by single and abundant insectivorous species, rather than by overall bird species richness. Forest proximity was important for enhancing the density of this key species, but did also promote bird species richness. Hence, our findings are both of economical as well as ecological interest because the conservation of nearby forest remnants will likely benefit human needs and biodiversity conservation alike.

Summary1. Biodiversity data are needed for conservation and management of tropical habitats, but the high diversity of these ecosystems makes comprehensive surveys prohibitively expensive and indicator taxa reflecting the biodiversity patterns of other taxa are frequently used. Few studies have produced the necessary comprehensive data sets to assess the quality of the indicator groups, however, and only one previous study has considered the monetary costs involved in sampling them. 2. We surveyed four plant groups (herbs, liverworts, trees, lianas) and eight animal groups (ants, canopy and dung beetles, birds, butterflies, bees, wasps and the parasitoids of the latter two) in 15 plots of 50 · 50 m 2 each, representing undisturbed rainforest and two types of cacao agroforest in Sulawesi, Indonesia. We calculated three biodiversity measures (a and b diversity; percentage of species indicative of habitat conditions), built simple and multiple regression models among species groups (single groups, combinations of 2-11 groups, averaged relative diversity of all 12 groups), and related these to three measures of survey cost (absolute costs and two approaches correcting for different sampling intensities). 3. Determination coefficients (R 2 values) of diversity patterns between single study groups were generally low (<0AE25), while the consideration of several study groups increased R 2 values to up to 0AE8 for combinations of four groups, and to almost 1AE0 for combinations of 11 groups. Survey costs varied 10-fold between study groups, but their cost-effectiveness (indicator potential versus monetary cost) varied strongly depending on the biodiversity aspect taken into account (a or b diversity, single or multiple groups, etc.). 4. Synthesis and applications. We found that increasing the number of taxa resulted in best overall biodiversity indication. We thus propose that the most cost-efficient approach to general tropical biodiversity inventories is to increase taxonomic coverage by selecting taxa with the lowest survey costs.

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.