Rainforest transformation reallocates energy from green to brown food webs.

Terrestrial animal biodiversity is increasingly being lost because of land-use change1,2. However, functional and energetic consequences aboveground and belowground and across trophic levels in megadiverse tropical ecosystems remain largely unknown. To fill this gap, we assessed changes in energy fluxes across 'green' aboveground (canopy arthropods and birds) and 'brown' belowground (soil arthropods and earthworms) animal food webs in tropical rainforests and plantations in Sumatra, Indonesia. Our results showed that most of the energy in rainforests is channelled to the belowground animal food web. Oil palm and rubber plantations had similar or, in the case of rubber agroforest, higher total animal energy fluxes compared to rainforest but the key energetic nodes were distinctly different: in rainforest more than 90% of the total animal energy flux was channelled by arthropods in soil and canopy, whereas in plantations more than 50% of the energy was allocated to annelids (earthworms). Land-use change led to a consistent decline in multitrophic energy flux aboveground, whereas belowground food webs responded with reduced energy flux to higher trophic levels, down to -90%, and with shifts from slow (fungal) to fast (bacterial) energy channels and from faeces production towards consumption of soil organic matter. This coincides with previously reported soil carbon stock depletion3. Here we show that well-documented animal biodiversity declines with tropical land-use change4-6 are associated with vast energetic and functional restructuring in food webs across aboveground and belowground ecosystem compartments.

Tree islands enhance biodiversity and functioning in oil palm landscapes.

In the United Nations Decade on Ecosystem Restoration1, large knowledge gaps persist on how to increase biodiversity and ecosystem functioning in cash crop-dominated tropical landscapes2. Here, we present findings from a large-scale, 5-year ecosystem restoration experiment in an oil palm landscape enriched with 52 tree islands, encompassing assessments of ten indicators of biodiversity and 19 indicators of ecosystem functioning. Overall, indicators of biodiversity and ecosystem functioning, as well as multidiversity and ecosystem multifunctionality, were higher in tree islands compared to conventionally managed oil palm. Larger tree islands led to larger gains in multidiversity through changes in vegetation structure. Furthermore, tree enrichment did not decrease landscape-scale oil palm yield. Our results demonstrate that enriching oil palm-dominated landscapes with tree islands is a promising ecological restoration strategy, yet should not replace the protection of remaining forests.

Balancing economic and ecological functions in smallholder and industrial oil palm plantations.

The expansion of the oil palm industry in Indonesia has improved livelihoods in rural communities, but comes at the cost of biodiversity and ecosystem degradation. Here, we investigated ways to balance ecological and economic outcomes of oil palm cultivation. We compared a wide range of production systems, including smallholder plantations, industrialized company estates, estates with improved agronomic management, and estates with native tree enrichment. Across all management types, we assessed multiple indicators of biodiversity, ecosystem functions, management, and landscape structure to identify factors that facilitate economic-ecological win-wins, using palm yields as measure of economic performance. Although, we found that yields in industrialized estates were, on average, twice as high as those in smallholder plantations, ecological indicators displayed substantial variability across systems, regardless of yield variations, highlighting potential for economic-ecological win-wins. Reducing management intensity (e.g., mechanical weeding instead of herbicide application) did not lower yields but improved ecological outcomes at moderate costs, making it a potential measure for balancing economic and ecological demands. Additionally, maintaining forest cover in the landscape generally enhanced local biodiversity and ecosystem functioning within plantations. Enriching plantations with native trees is also a promising strategy to increase ecological value without reducing productivity. Overall, we recommend closing yield gaps in smallholder cultivation through careful intensification, whereas conventional plantations could reduce management intensity without sacrificing yield. Our study highlights various pathways to reconcile the economics and ecology of palm oil production and identifies management practices for a more sustainable future of oil palm cultivation.

Land-use trajectories for sustainable land system transformations: Identifying leverage points in a global biodiversity hotspot.

Sustainable land-system transformations are necessary to avert biodiversity and climate collapse. However, it remains unclear where entry points for transformations exist in complex land systems. Here, we conceptualize land systems along land-use trajectories, which allows us to identify and evaluate leverage points, i.e., entry points on the trajectory where targeted interventions have particular leverage to influence land-use decisions. We apply this framework in the biodiversity hotspot Madagascar. In the northeast, smallholder agriculture results in a land-use trajectory originating in old-growth forests and spanning from forest fragments to shifting hill rice cultivation and vanilla agroforests. Integrating interdisciplinary empirical data on seven taxa, five ecosystem services, and three measures of agricultural productivity, we assess trade-offs and cobenefits of land-use decisions at three leverage points along the trajectory. These trade-offs and cobenefits differ between leverage points: Two leverage points are situated at the conversion of old-growth forests and forest fragments to shifting cultivation and agroforestry, resulting in considerable trade-offs, especially between endemic biodiversity and agricultural productivity. Here, interventions enabling smallholders to conserve forests are necessary. This is urgent since ongoing forest loss threatens to eliminate these leverage points due to path dependency. The third leverage point allows for the restoration of land under shifting cultivation through vanilla agroforests and offers cobenefits between restoration goals and agricultural productivity. The co-occurring leverage points highlight that conservation and restoration are simultaneously necessary to avert collapse of multifunctional mosaic landscapes. Methodologically, the framework highlights the importance of considering path dependency along trajectories to achieve sustainable land-system transformations.

Flower-bee versus pollen-bee metanetworks in fragmented landscapes.

Understanding the organization of mutualistic networks at multiple spatial scales is key to ensure biological conservation and functionality in human-modified ecosystems. Yet, how changing habitat and landscape features affect pollen-bee interaction networks is still poorly understood. Here, we analysed how bee-flower visitation and bee-pollen-transport interactions respond to habitat fragmentation at the local network and regional metanetwork scales, combining data from 29 fragments of calcareous grasslands, an endangered biodiversity hotspot in central Europe. We found that only 37% of the total unique pairwise species interactions occurred in both pollen-transport and flower visitation networks, whereas 28% and 35% were exclusive to pollen-transport and flower visitation networks, respectively. At local level, network specialization was higher in pollen-transport networks, and was negatively related to the diversity of land cover types in both network types. At metanetwork level, pollen transport data revealed that the proportion of single-fragment interactions increased with landscape diversity. Our results show that the specialization of calcareous grasslands' plant-pollinator networks decreases with landscape diversity, but network specialization is underestimated when only based on flower visitation information. Pollen transport data, more than flower visitation, and multi-scale analyses of metanetworks are fundamental for understanding plant-pollinator interactions in human-dominated landscapes.

Transformation scenarios towards multifunctional landscapes: A multi-criteria land-use allocation model applied to Jambi Province, Indonesia.

In tropical regions, shifting from forests and traditional agroforestry to intensive plantations generates conflicts between human welfare (farmers' demands and societal needs) and environmental protection. Achieving sustainability in this transformation will inevitably involve trade-offs between multiple ecological and socioeconomic functions. To address these trade-offs, our study used a new methodological approach allowing the identification of transformation scenarios, including theoretical landscape compositions that satisfy multiple ecological functions (i.e., structural complexity, microclimatic conditions, organic carbon in plant biomass, soil organic carbon and nutrient leaching losses), and farmers needs (i.e., labor and input requirements, total income to land, and return to land and labor) while accounting for the uncertain provision of these functions and having an actual potential for adoption by farmers. We combined a robust, multi-objective optimization approach with an iterative search algorithm allowing the identification of ecological and socioeconomic functions that best explain current land-use decisions. The model then optimized the theoretical land-use composition that satisfied multiple ecological and socioeconomic functions. Between these ends, we simulated transformation scenarios reflecting the transition from current land-use composition towards a normative multifunctional optimum. These transformation scenarios involve increasing the number of optimized socioeconomic or ecological functions, leading to higher functional richness (i.e., number of functions). We applied this method to smallholder farms in the Jambi Province, Indonesia, where traditional rubber agroforestry, rubber plantations, and oil palm plantations are the main land-use systems. Given the currently practiced land-use systems, our study revealed short-term returns to land as the principal factor in explaining current land-use decisions. Fostering an alternative composition that satisfies additional socioeconomic functions would require minor changes ("low-hanging fruits"). However, satisfying even a single ecological indicator (e.g., reduction of nutrient leaching losses) would demand substantial changes in the current land-use composition ("moonshot"). This would inevitably lead to a profit decline, underscoring the need for incentives if the societal goal is to establish multifunctional agricultural landscapes. With many oil palm plantations nearing the end of their production cycles in the Jambi province, there is a unique window of opportunity to transform agricultural landscapes.

Urbanization alters the spatiotemporal dynamics of plant-pollinator networks in a tropical megacity.

Urbanization is a major driver of biodiversity change but how it interacts with spatial and temporal gradients to influence the dynamics of plant-pollinator networks is poorly understood, especially in tropical urbanization hotspots. Here, we analysed the drivers of environmental, spatial and temporal turnover of plant-pollinator interactions (interaction ß-diversity) along an urbanization gradient in Bengaluru, a South Indian megacity. The compositional turnover of plant-pollinator interactions differed more between seasons and with local urbanization intensity than with spatial distance, suggesting that seasonality and environmental filtering were more important than dispersal limitation for explaining plant-pollinator interaction ß-diversity. Furthermore, urbanization amplified the seasonal dynamics of plant-pollinator interactions, with stronger temporal turnover in urban compared to rural sites, driven by greater turnover of native non-crop plant species (not managed by people). Our study demonstrates that environmental, spatial and temporal gradients interact to shape the dynamics of plant-pollinator networks and urbanization can strongly amplify these dynamics.

Tree identity and canopy openness mediate oil palm biodiversity enrichment effects on insect herbivory and pollination.

As the extent of oil palm (Elaeis guineensis) cultivation has expanded at the expense of tropical rainforests, enriching conventional large-scale oil palm plantations with native trees has been proposed as a strategy for restoring biodiversity and ecosystem function. However, how tree enrichment affects insect-mediated ecosystem functions is unknown. We investigated impacts on insect herbivory and pollination in the fourth year of a plantation-scale, long-term oil palm biodiversity enrichment experiment in Jambi, Sumatra, Indonesia. Within 48 plots systematically varying in size (25-1600 m2 ) and planted tree species richness (one to six species), we collected response data on vegetation structure, understory insect abundances, and pollinator and herbivore activity on chili plants (Capsicum annuum), which served as indicators of insect-mediated ecosystem functions. We examined the independent effects of plot size, tree species richness, and tree identity on these response variables, using the linear model for random partitions design. The experimental treatments were most associated with vegetation structure: tree identity mattered, as the species Peronema canescens strongly decreased (by approximately one standard deviation) both canopy openness and understory vegetation cover; whereas tree richness only decreased understory flower density. Further, the smallest plots had the lowest understory flower density and richness, presumably because of lower light availability and colonization rates, respectively. Enrichment influenced herbivorous insects and natural enemies in the understory to a lesser extent: both groups had higher abundances in plots with two enrichment species planted, possibly because higher associated tree mortality created more habitat, while herbivores decreased with increasing tree species richness, in line with the resource concentration hypothesis. Linking relationships in structural equation models showed that the negative association between P. canescens and understory vegetation cover was mediated through canopy openness. Likewise, canopy openness mediated increases in herbivore and pollinator insect abundances. Higher pollinator visitation increased phytometer yield, while impacts of insect herbivores on yield were not apparent. Our results demonstrate that even at an early stage, different levels of ecological restoration influence insect-mediated ecosystem functions, mainly through canopy openness. These findings suggest that maintaining some canopy gaps while enrichment plots develop may be beneficial for increasing habitat heterogeneity and insect-mediated ecosystem functions.

Functional diversity of farmland bees across rural-urban landscapes in a tropical megacity.

Urbanization poses a major threat to biodiversity and food security, as expanding cities, especially in the Global South, increasingly compete with natural and agricultural lands. However, the impact of urban expansion on agricultural biodiversity in tropical regions is overlooked. Here we assess how urbanization affects the functional response of farmland bees, the most important pollinators for crop production. We sampled bees across three seasons in 36 conventional vegetable-producing farms spread along an urbanization gradient in Bengaluru, an Indian megacity. We investigated how landscape and local environmental drivers affected different functional traits (sociality, nesting behavior, body size, and specialization) and functional diversity (functional dispersion) of bee communities. We found that the functional responses to urbanization were trait specific with more positive than negative effects of gray area (sealed surfaces and buildings) on species richness, functional diversity, and abundance of most functional groups. As expected, larger, solitary, cavity-nesting, and, surprisingly, specialist bees benefited from urbanization. In contrast to temperate cities, the abundance of ground nesters increased in urban areas, presumably because larger patches of bare soil were still available beside roads and buildings. However, overall bee abundance and the abundance of social bees (85% of all bees) decreased with urbanization, threatening crop pollination. Crop diversity promotes taxonomic and functional diversity of bee communities. Locally, flower resources promote the abundance of all functional groups, and natural vegetation can maintain diverse pollinator communities throughout the year, especially during the noncropping season. However, exotic plants decrease functional diversity and bee specialization. To safeguard bees and their pollination services in urban farms, we recommend (1) preserving seminatural vegetation (hedges) around cropping fields to provide nesting opportunities for aboveground nesters, (2) promoting farm-level crop diversification of beneficial crops (e.g., pulses, vegetables, and spices), (3) maintaining native natural vegetation along field margins, and (4) controlling and removing invasive exotic plants that disrupt native plant-pollinator interactions. Overall, our results suggest that urban agriculture can maintain functionally diverse bee communities and, if managed in a sustainable manner, be used to develop win-win solutions for biodiversity conservation of pollinators and food security in and around cities.

Floral resource diversification promotes solitary bee reproduction and may offset insecticide effects - evidence from a semi-field experiment.

Pollinator declines in agricultural landscapes are driven by multiple stressors, but potential interactions of these remain poorly studied. Using a highly replicated semi-field study with 56 mesocosms of varying wild plant diversity (2-16 species) and oilseed rape treated with a neonicotinoid, we tested the interacting effects of resource diversity and insecticides on reproduction of a solitary wild bee. Compared to mesocosms with oilseed rape monocultures, availability of resources from wild plants complementing oilseed rape doubled brood cell production. In addition, bee reproduction increased due to plant diversity and identity effects. Exposure to neonicotinoid-treated oilseed rape reduced bee larval to adult development by 69%, but only in mesocosms with oilseed rape monocultures. Availability of complementary flower resources can thus offset negative effects of neonicotinoid-treated oilseed rape on wild bee reproduction. Policy should encourage the implementation of diverse floral resources mitigating negative effects of crop monocultures and insecticides, thereby sustaining solitary bee populations in agricultural landscapes.

A plant-pollinator metanetwork along a habitat fragmentation gradient.

To understand how plant-pollinator interactions respond to habitat fragmentation, we need novel approaches that can capture properties that emerge at broad scales, where multiple communities engage in metanetworks. Here we studied plant-pollinator interactions over 2 years on 29 calcareous grassland fragments selected along independent gradients of habitat size and surrounding landscape diversity of cover types. We associated network centrality of plant-pollinator interactions and grassland fragments with their ecological and landscape traits, respectively. Interactions involving habitat specialist plants and large-bodied pollinators were the most central, implying that species with these traits form the metanetwork core. Large fragments embedded in landscapes with high land cover diversity exhibited the highest centrality; however, small fragments harboured many unique interactions not found on larger fragments. Intensively managed landscapes have reached a point in which all remaining fragments matter, meaning that losing any further areas may vanish unique interactions with unknown consequences for ecosystem functioning.

Taxonomic and functional homogenization of farmland birds along an urbanization gradient in a tropical megacity.

Urbanization is a major driver of land use change and biodiversity decline. While most of the ongoing and future urbanization hotspots are located in the Global South, the impact of urban expansion on agricultural biodiversity and associated functions and services in these regions has widely been neglected. Additionally, most studies assess biodiversity responses at local scale (α-diversity), however, ecosystem functioning is strongly determined by compositional and functional turnover of communities (ß-diversity) at regional scales. We investigated taxonomic and functional ß-diversity of farmland birds across three seasons on 36 vegetable farms spread along a continuous urbanization gradient in Bangalore, a South Indian megacity. Increasing amount of grey area in the farm surroundings was the dominant driver affecting ß-diversity and resulting in taxonomic and functional homogenization of farmland bird communities. Functional diversity losses were higher than expected from species declines (i.e., urbanization acts as an environmental filter), with particular losses of functionally important groups such as insectivores of crop pests. Moreover, urbanization reduced functional redundancy of bird communities, which may further weaken ecosystems resilience to future perturbations. Our study underscores urbanization as a major driver of taxonomic and functional homogenization of species communities in agricultural systems, potentially threatening crucial ecosystem services for food production.

Partitioning wild bee and hoverfly contributions to plant-pollinator network structure in fragmented habitats.

The risk of ecosystem function degradation with biodiversity loss has emerged as a major scientific concern in recent years. Possible relationships between taxonomic diversity and magnitude and stability of ecosystem processes build upon species' functional characteristics, which determine both susceptibility to environmental change and contribution to ecosystem properties. The functional diversity within communities thus provides a potential buffer against environmental disturbance, especially for properties emerging from interactions among species. In complex plant-pollinator networks, distantly related taxa spanning a great trait diversity shape network architecture. Here, we address the question of whether network properties are maintained after habitat loss by complementary contributions of phylogenetically distant pollinator taxa. We quantified contributions of wild bees and hoverflies to network structure (connectance, network specialization, specialization asymmetry) in 32 calcareous grassland fragments varying in size. Although hoverflies are often regarded less susceptible to environmental change than wild bees, species richness of both taxa was similarly affected by habitat loss. The associated loss of 80% of interactions resulted in small and tightly connected networks, which was more strongly attributed to wild bee loss than hoverfly loss. Networks in small fragments were less specialized due to equivalent losses of species and interactions in both pollinators and plants. Because wild bee and hoverfly loss contributed similarly to declining network specialization, we conclude that trait diversity among distantly related pollinators does not necessarily provide insurance against functional homogenization during community disassembly following habitat loss.

Autonomous sound recording outperforms human observation for sampling birds: a systematic map and user guide.

Autonomous sound recording techniques have gained considerable traction in the last decade, but the question remains whether they can replace human observation surveys to sample sonant animals. For birds in particular, survey methods have been tested extensively using point counts and sound recording surveys. Here, we review the latest evidence for this taxon within the frame of a systematic map. We compare sampling effectiveness of these two survey methods, the output variables they produce, and their practicality. When assessed against the standard of point counts, autonomous sound recording proves to be a powerful tool that samples at least as many species. This technology can monitor birds in an exhaustive, standardized, and verifiable way. Moreover, sound recorders give access to entire soundscapes from which new data types can be derived (vocal activity, acoustic indices). Variables such as abundance, density, occupancy, or species richness can be obtained to yield data sets that are comparable to and compatible with point counts. Finally, autonomous sound recorders allow investigations at high temporal and spatial resolution and coverage, which are more cost effective and cannot be achieved by human observations alone, even though small-scale studies might be more cost effective when carried out with point counts. Sound recorders can be deployed in many places, they are more scalable and reliable, making them the better choice for bird surveys in an increasingly data-driven time. We provide an overview of currently available recorders and discuss their specifications to guide future study designs.

Trophic level, successional age and trait matching determine specialization of deadwood-based interaction networks of saproxylic beetles.

The specialization of ecological networks provides important insights into possible consequences of biodiversity loss for ecosystem functioning. However, mostly mutualistic and antagonistic interactions of living organisms have been studied, whereas detritivore networks and their successional changes are largely unexplored. We studied the interactions of saproxylic (deadwood-dependent) beetles with their dead host trees. In a large-scale experiment, 764 logs of 13 tree species were exposed to analyse network structure of three trophic groups of saproxylic beetles over 3 successional years. We found remarkably high specialization of deadwood-feeding xylophages and lower specialization of fungivorous and predatory species. During deadwood succession, community composition, network specialization and network robustness changed differently for the functional groups. To reveal potential drivers of network specialization, we linked species' functional traits to their network roles, and tested for trait matching between plant (i.e. chemical compounds) and beetle (i.e. body size) traits. We found that both plant and animal traits are major drivers of species specialization, and that trait matching can be more important in explaining interactions than neutral processes reflecting species abundance distributions. High network specialization in the early successional stage and decreasing network robustness during succession indicate vulnerability of detritivore networks to reduced tree species diversity and beetle extinctions, with unknown consequences for wood decomposition and nutrient cycling.

Insectivorous birds disrupt biological control of cereal aphids.

Insect-feeding birds may interfere with trophic interactions in plant-insect food webs, which may be particularly important in agroecosystems. Here, we studied how Eurasian Tree Sparrows (Passer montanus) affect aphids and their predators in cereal fields using bird exclusion experiments. The Tree Sparrows fed their nestlings with aphid antagonists. Hoverflies and ladybird beetles accounted for 77% of the food for the nestlings during peak aphid density. When birds were excluded, densities of hoverfly larvae, which were the most abundant aphid predator group in the cereals, were 4% higher in wheat and 45% higher in oat, while aphid densities were 24% lower in wheat and 26% lower in oat. The demonstrated disruption of biological control by mesopredators through bird predation may be a common phenomenon in cropping systems characterized by small-sized and abundant pest species. Management of biotic interactions such as biological control needs a broad food-web perspective, even in simplified agroecosystems.

The role of ants, birds and bats for ecosystem functions and yield in oil palm plantations.

One of the world's most important and rapidly expanding crops, oil palm, is associated with low levels of biodiversity. Changes in predator communities might alter ecosystem services and subsequently sustainable management but these links have received little attention to date. Here, for the first time, we manipulated ant and flying vertebrate (birds and bats) access to oil palms in six smallholder plantations in Sumatra (Indonesia) and measured effects on arthropod communities, related ecosystem functions (herbivory, predation, decomposition and pollination) and crop yield. Arthropod predators increased in response to reductions in ant and bird access, but the overall effect of experimental manipulations on ecosystem functions was minimal. Similarly, effects on yield were not significant. We conclude that ecosystem functions and productivity in oil palm are, under current levels of low pest pressure and large pollinator populations, robust to large reductions of major predators.

Mixing on- and off-field measures for biodiversity conservation.

The continuing biodiversity losses through agricultural expansion and intensification are dramatic. We argue that a mix of on- and off-field measures is needed, overcoming the false dichotomy of the land sharing-sparing debate. Protected land is essential for global biodiversity, while spillover between farmed and natural land is key to reducing species extinctions. This is particularly effective in landscapes with small and diversified fields. Focusing only on protected land fails to conserve a wealth of species, which often provide major ecosystem services such as pest control, pollination, and cultural benefits. On-field measures must minimise yield losses to prevent increased demand for food imports from biodiversity-rich regions, requiring enforcement of high social-ecological land-use standards to ensure a good life for all.

Joint environmental and social benefits from diversified agriculture.

Agricultural simplification continues to expand at the expense of more diverse forms of agriculture. This simplification, for example, in the form of intensively managed monocultures, poses a risk to keeping the world within safe and just Earth system boundaries. Here, we estimated how agricultural diversification simultaneously affects social and environmental outcomes. Drawing from 24 studies in 11 countries across 2655 farms, we show how five diversification strategies focusing on livestock, crops, soils, noncrop plantings, and water conservation benefit social (e.g., human well-being, yields, and food security) and environmental (e.g., biodiversity, ecosystem services, and reduced environmental externalities) outcomes. We found that applying multiple diversification strategies creates more positive outcomes than individual management strategies alone. To realize these benefits, well-designed policies are needed to incentivize the adoption of multiple diversification strategies in unison.

Additive effects of exotic plant abundance and land-use intensity on plant-pollinator interactions.

The continuing spread of exotic plants and increasing human land-use are two major drivers of global change threatening ecosystems, species and their interactions. Separate effects of these two drivers on plant-pollinator interactions have been thoroughly studied, but we still lack an understanding of combined and potential interactive effects. In a subtropical South African landscape, we studied 17 plant-pollinator networks along two gradients of relative abundance of exotics and land-use intensity. In general, pollinator visitation rates were lower on exotic plants than on native ones. Surprisingly, while visitation rates on native plants increased with relative abundance of exotics and land-use intensity, pollinator visitation on exotic plants decreased along the same gradients. There was a decrease in the specialization of plants on pollinators and vice versa with both drivers, regardless of plant origin. Decreases in pollinator specialization thereby seemed to be mediated by a species turnover towards habitat generalists. However, contrary to expectations, we detected no interactive effects between the two drivers. Our results suggest that exotic plants and land-use promote generalist plants and pollinators, while negatively affecting specialized plant-pollinator interactions. Weak integration and high specialization of exotic plants may have prevented interactive effects between exotic plants and land-use. Still, the additive effects of exotic plants and land-use on specialized plant-pollinator interactions would have been overlooked in a single-factor study. We therefore highlight the need to consider multiple drivers of global change in ecological research and conservation management.