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Tropical forests are threatened by degradation and deforestation but the consequences for these ecosystems are poorly understood, particularly at the landscape scale. We present the most extensive ecosystem analysis to date of the impacts of logging and conversion of tropical forest to oil palm from a large-scale study in Borneo, synthesizing responses from 79 variables categorized into four hierarchical ecological ‘levels’: 1) structure and environment, 2) species traits, 3) biodiversity and 4) ecosystem functions. Variables at the lowest levels that were directly impacted by the physical processes of timber extraction, such as soil characteristics, were sensitive to even moderate amounts of logging, whereas biodiversity and ecosystem functions proved remarkably resilient to logging in many cases, but were more affected by conversion to oil palm plantation.One-Sentence SummaryLogging tropical forest mostly impacts structure while biodiversity and functions are more vulnerable to habitat conversion

Agricultural expansion is the primary driver of ecological degradation across the tropics. Set-asides—uncultivated parts of agricultural landscapes, often on steep slopes and alongside rivers—may alleviate environmental impacts but can reduce the area cultivated. Here we model an approach to configuring set-asides aimed at optimizing ecological outcomes (biodiversity, above-ground carbon storage and nutrient cycling) without reducing net cultivation area. We compare set-asides in an oil palm landscape where all plantations adopt the same configuration (‘uniform’ approach) with a scenario where there can be variation in configuration among plantations (‘variable’ approach). We find that all set-aside configurations support substantial ecological values but that the best strategies involve set-asides, particularly alongside rivers, that are spatially targeted and variable among plantations. This ‘variable’ approach can increase ecological outcomes twofold over the ‘uniform’ approach without reducing net cultivation area. Our findings underscore the potential importance of well-planned set-asides for enhancing agricultural sustainability.

Wallacea-the meeting point between the Asian and Australian fauna-is one of the world's largest centers of endemism. Twenty-three million years of complex geological history have given rise to a living laboratory for the study of evolution and biodiversity, highly vulnerable to anthropogenic pressures. In the present article, we review the historic and contemporary processes shaping Wallacea's biodiversity and explore ways to conserve its unique ecosystems. Although remoteness has spared many Wallacean islands from the severe overexploitation that characterizes many tropical regions, industrial-scale expansion of agriculture, mining, aquaculture and fisheries is damaging terrestrial and aquatic ecosystems, denuding endemics from communities, and threatening a long-term legacy of impoverished human populations. An impending biodiversity catastrophe demands collaborative actions to improve community-based management, minimize environmental impacts, monitor threatened species, and reduce wildlife trade. Securing a positive future for Wallacea's imperiled ecosystems requires a fundamental shift away from managing marine and terrestrial realms independently.

Anthropogenic-driven species extinctions are radically changing the biosphere. Biological communities may become increasingly similar to or dissimilar from one another via the processes of biotic homogenisation or heterogenisation. A key question is how the conversion of native forests to agriculture may influence these processes by driving changes in the occurrence patterns of restricted-range endemic species versus wide-ranging generalists. We examined biotic homogenisation and heterogenisation in bird communities on the Southeast Asian islands of Borneo, Sulawesi, Seram, Buru, Talaud and Sangihe. Each island is characterised by high levels of avian endemism, and unique spatial configuration of forest conversion to agriculture. Forest conversion to agriculture influenced the patterns of biotic homogenisation on five islands. Bird communities became increasingly dissimilar to forest reference communities relative to localised patterns of deforestation. Turnover led to species with larger global range-sizes dominating communities at the expense of island endemics and ecological specialists. Within islands, forest conversion did not result in clear changes to β-diversity, whereas between-island communities became increasingly similar with greater deforestation, implying that patterns of forest conversion profoundly affect biotic homogenisation. Our findings elucidate how continued conversion of forests is causing the replacement of endemic species by a small cohort of shared ubiquitous species with potentially strong negative consequences for ecosystem functioning and resilience. Halting reorganisation of the biosphere via the loss of range-restricted species and spread of wide-ranged generalists will require improved efforts to reduce the impacts of deforestation particularly in regions with high endemism.

Agricultural expansion drives biodiversity decline in forested tropical regions. Consequently, it is important to understand the conservation value of remnant forest in production landscapes. In a tropical landscape dominated by oil palm we characterized faunal communities across eight taxa occurring within riparian forest buffers, which are legally protected alongside rivers, and compared them to nearby recovering logged forest. Buffer width was the main predictor of species richness and abundance, with widths of 40-100 m on each side of the river supporting broadly equivalent levels of biodiversity to logged forest. However, width responses varied markedly among taxa, and buffers often lacked forest-dependent species. Much wider buffers than are currently mandated are needed to safeguard most species. The largest biodiversity gains are achieved by increasing relatively narrow buffers. To provide optimal conservation outcomes in tropical production landscapes we encourage policymakers to prescribe width requirements for key taxa and different landscape contexts., Newton-Ungku Omar Fund (grants 216433953, 537134717) – delivered by the British Council and funded by the UK Department for Business, Energy and Industrial Strategy and the Malaysian Industry-Government Group for High Technology – as well as the UK Natural Environment Research Council (NE/K016407/1, NE/K016261/1; https://lombok.nerc-hmtf.info/). MJS was supported by a Research Leadership Award from the Leverhulme Trust.

Avid birdwatchers, or ‘twitchers’, expend a considerable amount of money and time pursuing viewing experiences of rare or vagrant species. By vagrant species, we mean a species found outside its normal range/distribution. To enhance our understanding of this form of behaviour, we present results from a U.K. survey of twitchers. First, we examine the relationship between cost and rarity based on actual viewing experiences. Our statistical results reveal that the relationship between cost and rarity is positive and very inelastic. Second, we present results from a hypothetical Best-Worst Scaling exercise examining aspects of species rarity. We find that rarity is a more nuanced construct than simply the frequency with which a vagrant species has appeared. Our results provide insights into the meaning of rarity, as well as the economic value attach to it and why.

The Wallacea biogeographic region of Sulawesi, the Moluccas and Lesser Sunda is globally renowned for exceptional endemism, but is currently emerging as a development frontier in Indonesia. We assessed patterns and drivers of forest loss and fragmentation across the region, and used dynamic deforestation models to project future deforestation to 2053. Up to 10 231 km2 was deforested between 2000 and 2018, and a further 49 570 km2 is expected to be lost by 2053, with annual deforestation rates ranging between 0.09% and 2.17% in different sub-regions (average: 1.23%). Key biodiversity areas (priority sites for endemic and threatened biodiversity) are particularly vulnerable to deforestation if they are small, coastal and unprotected. Sub-regional variation in deforestation patterns and drivers must be acknowledged if conservation interventions are to be targeted and effective. We provide a valuable baseline from which to monitor Wallacea’s new development course, as Indonesia undergoes profound policy changes that will provide both challenges and opportunities for environmental governance and conservation.

Measuring the multidimensional diversity properties of a community is of great importance for ecologists, conservationists and stakeholders. Diversity profiles, a plotted series of Hill numbers, simultaneously capture the common diversity indices. However, diversity metrics require information on species abundance, often relying on raw count data without accounting for imperfect and varying detection. Hierarchical occupancy models account for variation in detectability, and Hill numbers have been expanded to allow estimation based on occupancy probability. But the ability of occupancy-based diversity profiles to reproduce patterns in abundance-based diversity has not been investigated. Here, we fit community occupancy models to simulated animal communities to explore how well occupancy-based diversity profiles reflect patterns in true abundance-based diversity. Because we expect occupancy-based diversity to be overestimated, we further tested a occupancy thresholding approach to reduce potential biases in the estimated diversity profiles. Finally, we use empirical bird community data to present how the framework can be extended to consider species similarity. The simulation study showed that occupancy-based diversity profiles produced among-community patterns in diversity similar to true abundance diversity profiles, although within-community diversity was generally overestimated. Applying an occupancy threshold reduced positive bias, but resulted in negative bias in richness estimates and slightly reduced the ability to reproduce true differences among the simulated communities; thus, we do not recommend application of this threshold. Application of our approach to a large bird dataset indicated differential species diversity patterns in communities of different habitat types. Accounting for phylogenetic and ecological similarities between species reduced variability in diversity among habitats. Our framework allows investigating the complexity of diversity from species detection data, while accounting for imperfect and varying detection probabilities, as well as species similarities. Visualizing results in the form of diversity profiles facilitates comparison of diversity between sites or across time. The approach offers opportunities for further development, for example by using local abundances estimated using the Royle-Nichols or N-mixture models and further exploration of thresholding methods. In spite of some challenges, occupancy-based diversity profiles are useful for studying and monitoring patterns in biodiversity.

Agricultural expansion across the tropics is the primary driver of biodiversity declines and ecosystem service degradation. However, efforts to mitigate these negative impacts may reduce commodity production. We quantify trade-offs between oil palm cultivation and ecological outcomes (biodiversity, above-ground carbon storage and dung nutrient cycling) across different potential set-aside (uncultivated areas in agricultural landscapes) strategies. We show that all set-aside configurations yield substantial gains in ecological outcomes. The best strategy involves spatially targeted riparian reserves, such as those used in oil palm certification schemes, where species occurrence can be doubled without reducing overall cultivation area. Adopting this strategy throughout the 8 million hectares of plantations in Borneo would lead to extensive improvements in ecological outcomes without losses to production area, and consequently, enhancing agricultural sustainability.

Approaches to characterise and monitor biodiversity based on the sound signals of ecosystems have become popular in landscape ecology and biodiversity conservation. However, to date, validation studies of how well acoustic indices reflect observed biodiversity patterns have often relied on low levels of either spatial or temporal replication, while focussing on habitats with similar underlying anthropological and geophysical sound characteristics. For acoustic indices to be broadly applicable to biodiversity monitoring, their capacity to measure the ecological facets of soundscapes must be robust to these potential sources of bias. Using two contrasting recording approaches, we examined the efficacy of four commonly used acoustic indices to reflect patterns of observed bird species richness across a tropical forest degradation gradient in Northeast Borneo. The gradient comprised intact and logged forests, riparian forests, remnants, and oil palm plantations, thus providing a highly variable anthrophonic and geophonic soundscape. We compared the degree to which acoustic indices derived from automated versus point count recording methods detected variation in inter-habitat species richness, as well as their capacity to capture changes in species diversity as a consequence of forest degradation quantified by high-resolution LiDAR derived forest canopy heights. We found Acoustic Diversity Index was associated with forest canopy height as measured by both automated recorders and recordings from point counts, whereas the association between canopy height and Acoustic Complexity Index was only detected using point count recordings. For both types of recordings, Acoustic Complexity Index exhibited the strongest relationship with observed bird richness in old growth and logged forest, whereas Acoustic Diversity was not linked, suggesting avian richness does not drive its association with canopy height. No acoustic indices were associated with observed bird richness in oil palm riparian areas. Our findings underscore the potential utility of soundscape approaches to characterise biodiversity patterns in degraded tropical landscapes, and may be used as a proxy for human inventories of bird communities. However, we also show that for acoustic indices to be effective on landscape-wide studies of environmental gradients, adequate spatial replication is required, and care must be taken to control for non-target elements of soundscapes in different habitats.

The expansion of oil palm agriculture has contributed to biodiversity loss in Southeast Asia and elsewhere in the tropics. Riparian reserves (areas of native forest along waterways) have the potential to maintain forest biodiversity and associated ecological processes within these agricultural landscapes. Using acoustic sampling, we investigated the value of riparian reserves for insectivorous bats in oil palm plantations in Sabah, Malaysian Borneo. We compared general bat activity, foraging activity, and species occupancy between riparian areas in forest and riparian reserves in oil palm plantations. Overall bat activity varied little between riparian reserves in oil palm and riparian forest. Rather, activity was greatest in areas with a high forest canopy, irrespective of how much forest was available within or outside the riparian reserve. Bat foraging activity, as well as the occupancy of two species, was greatest in the forest sites, and while bats were detected in the oil palm riparian reserves, both foraging and occupancy were more associated with topographic ruggedness than forest amount or height. Our results indicate that habitat structure within riparian reserves may be more important than reserve size for supporting insectivorous bat diversity within oil palm landscapes. These findings provide important insights into the extent of the ecological benefits provided by conservation set-asides in forest-agricultural landscapes in the tropics.

Zero‐deforestation commitments seek to decouple agricultural production and forest loss to improve prospects for biodiversity. However, the effectiveness of methods designed to meet these commitments is poorly understood. In a highly fragmented tropical landscape dominated by oil palm, we tested the capacity for the High Carbon Stock (HCS) Approach to prioritize forest remnants that sustain mammal diversity. Patches afforded high priority by HCS protocols (100 ha core area) provided important refuges for IUCN‐threatened species and megafauna. However, patch‐scale HCS area recommendations conserved only 35% of the mammal community. At least 3,000 ha would be required to retain intact mammal assemblages, with nearly 10 times this area needed if hunting pressure was high. While current HCS protocols will safeguard patches capable of sustaining biodiversity, highly fragmented tropical landscapes typical of zero‐deforestation pledges will require thinking beyond the patch toward strategically configured forest remnants at the landscape level and enforcing strict controls on hunting.

1. Conversion of forest to oil palm agriculture is a significant and continuing threat to tropical biodiversity. Despite this, little is known about the value of riparian reserves in oil palm and how these conservation set-asides might best be managed to maintain biodiversity.\ud \ud 2. We quantified bird community structure of 20 sites in an oil palm-forest mosaic in Sabah, Malaysia using 6104 detections from 867 point counts. Sites included oil palm riparian reserves of various vegetation quality and reserve widths, which were compared to oil palm streams without a riparian reserve, riparian areas in continuous logged forest and non-riparian logged forest control sites.\ud \ud 3. Riparian reserves, oil palm waterways, continuous riparian forest and non-riparian control forest sites supported distinct avifaunal communities. Generalised linear models showed riparian reserve width, forest quality and the amount of surrounding forest cover to be the strongest predictors of bird species richness. For forest dependent species, each of these predictors had stronger effect size when compared with all species. On average, reserves held 31% of all species and 30% of forest specialists, whereas continuous riparian forests averaged 32% of all species but 38% of forest species.\ud \ud 4. Riparian reserves with >40 m of natural vegetation on each bank supported similar bird diversity to riparian habitats found in continuous forest. However, to support equivalent numbers of forest-dependent species and species of conservation concern, reserves would need to be at least 100 m wide on each bank. The highest numbers of species were found in riparian forest with above-ground carbon densities exceeding 75 tC ha-1, indicating the importance of habitat quality, as well as width, in supporting riparian bird communities.\ud \ud 5. Synthesis and applications. If designed and protected appropriately, riparian reserves support diverse bird communities including many species of conservation concern in oil palm estates. This can be achieved by designating large reserves (80-200 m total width), but to maximize species numbers forest disturbance should also be minimised prior to conversion as well as during plantation operations.

Selective logging is a major driver of rainforest degradation across the tropics. Two competing logging strategies are proposed to meet timber demands with the least impact on biodiversity: land sharing, which combines timber extraction with biodiversity protection across the concession; and land sparing, in which higher intensity logging is combined with the protection of intact primary forest reserves. We evaluate these strategies by comparing the abundances and species richness of birds, dung beetles and ants in Borneo, using a protocol that allows us to control for both timber yield and net profit across strategies. Within each taxonomic group, more species had higher abundances with land-sparing than land-sharing logging, and this translated into significantly higher species richness within land-sparing concessions. Our results are similar when focusing only on species found in primary forest and restricted in range to Sundaland, and they are independent of the scale of sampling. For each taxonomic group, land-sparing logging was the most promising strategy for maximizing the biological value of logging operations.

Effects of logging on species composition in tropical rainforests are well known but may fail to reveal key changes in species interactions. We used nitrogen stable-isotope analysis of 73 species of understory birds to quantify trophic responses to repeated intensive logging of rainforest in northern Borneo and to test 4 hypotheses: logging has significant effects on trophic positions and trophic-niche widths of species, and the persistence of species in degraded forest is related to their trophic positions and trophic-niche widths in primary forest. Species fed from higher up the food chain and had narrower trophic-niche widths in degraded forest. Species with narrow trophic-niche widths in primary forest were less likely to persist after logging, a result that indicates a higher vulnerability of dietary specialists to local extinction following habitat disturbance. Persistence of species in degraded forest was not related to a species' trophic position. These results indicate changes in trophic organization that were not apparent from changes in species composition and highlight the importance of focusing on trophic flexibility over the prevailing emphasis on membership of static feeding guilds. Our results thus support the notion that alterations to trophic organization and interactions within tropical forests may be a pervasive and functionally important hidden effect of forest degradation.

Effects of logging on species composition in tropical rainforests are well known but may fail to reveal key changes in species interactions. We used nitrogen stable-isotope analysis of 73 species of understory birds to quantify trophic responses to repeated intensive logging of rainforest in northern Borneo and to test 4 hypotheses: logging has significant effects on trophic positions and trophic-niche widths of species, and the persistence of species in degraded forest is related to their trophic positions and trophic-niche widths in primary forest. Species fed from higher up the food chain and had narrower trophic-niche widths in degraded forest. Species with narrow trophic-niche widths in primary forest were less likely to persist after logging, a result that indicates a higher vulnerability of dietary specialists to local extinction following habitat disturbance. Persistence of species in degraded forest was not related to a species' trophic position. These results indicate changes in trophic organization that were not apparent from changes in species composition and highlight the importance of focusing on trophic flexibility over the prevailing emphasis on membership of static feeding guilds. Our results thus support the notion that alterations to trophic organization and interactions within tropical forests may be a pervasive and functionally important hidden effect of forest degradation.