The future of hyperdiverse tropical ecosystems.

The tropics contain the overwhelming majority of Earth's biodiversity: their terrestrial, freshwater and marine ecosystems hold more than three-quarters of all species, including almost all shallow-water corals and over 90% of terrestrial birds. However, tropical ecosystems are also subject to pervasive and interacting stressors, such as deforestation, overfishing and climate change, and they are set within a socio-economic context that includes growing pressure from an increasingly globalized world, larger and more affluent tropical populations, and weak governance and response capacities. Concerted local, national and international actions are urgently required to prevent a collapse of tropical biodiversity.

Anthropogenic disturbance in tropical forests can double biodiversity loss from deforestation.

Concerted political attention has focused on reducing deforestation, and this remains the cornerstone of most biodiversity conservation strategies. However, maintaining forest cover may not reduce anthropogenic forest disturbances, which are rarely considered in conservation programmes. These disturbances occur both within forests, including selective logging and wildfires, and at the landscape level, through edge, area and isolation effects. Until now, the combined effect of anthropogenic disturbance on the conservation value of remnant primary forests has remained unknown, making it impossible to assess the relative importance of forest disturbance and forest loss. Here we address these knowledge gaps using a large data set of plants, birds and dung beetles (1,538, 460 and 156 species, respectively) sampled in 36 catchments in the Brazilian state of Pará. Catchments retaining more than 69­80% forest cover lost more conservation value from disturbance than from forest loss. For example, a 20% loss of primary forest, the maximum level of deforestation allowed on Amazonian properties under Brazil's Forest Code, resulted in a 39­54% loss of conservation value: 96­171% more than expected without considering disturbance effects. We extrapolated the disturbance-mediated loss of conservation value throughout Pará, which covers 25% of the Brazilian Amazon. Although disturbed forests retained considerable conservation value compared with deforested areas, the toll of disturbance outside Pará's strictly protected areas is equivalent to the loss of 92,000­139,000 km2 of primary forest. Even this lowest estimate is greater than the area deforested across the entire Brazilian Amazon between 2006 and 2015 (ref. 10). Species distribution models showed that both landscape and within-forest disturbances contributed to biodiversity loss, with the greatest negative effects on species of high conservation and functional value. These results demonstrate an urgent need for policy interventions that go beyond the maintenance of forest cover to safeguard the hyper-diversity of tropical forest ecosystems.

Linking global drivers of agricultural trade to on-the-ground impacts on biodiversity.

Consumption of globally traded agricultural commodities like soy and palm oil is one of the primary causes of deforestation and biodiversity loss in some of the world's most species-rich ecosystems. However, the complexity of global supply chains has confounded efforts to reduce impacts. Companies and governments with sustainability commitments struggle to understand their own sourcing patterns, while the activities of more unscrupulous actors are conveniently masked by the opacity of global trade. We combine state-of-the-art material flow, economic trade, and biodiversity impact models to produce an innovative approach for understanding the impacts of trade on biodiversity loss and the roles of remote markets and actors. We do this for the production of soy in the Brazilian Cerrado, home to more than 5% of the world´s species. Distinct sourcing patterns of consumer countries and trading companies result in substantially different impacts on endemic species. Connections between individual buyers and specific hot spots explain the disproportionate impacts of some actors on endemic species and individual threatened species, such as the particular impact of European Union consumers on the recent habitat losses for the iconic giant anteater (Myrmecophaga tridactyla). In making these linkages explicit, our approach enables commodity buyers and investors to target their efforts much more closely to improve the sustainability of their supply chains in their sourcing regions while also transforming our ability to monitor the impact of such commitments over time.

A Commodity Supply Mix for More Regionalized Life Cycle Assessments.

Supply chain information is invaluable to further regionalize product life cycle assessments (LCAs), but detailed information linking production and consumption centers is not always available. We introduce the commodity supply mix (CSM) defined as the trade-volume-weighted average representing the combined geographic areas for the production of a commodity exported to a given market with the goal of (1) enhancing the relevance of inventory and impact regionalization and (2) allocating these impacts to specific markets. We apply the CSM to the Brazilian soybean supply chain mapped by Trase to obtain the mix of ecoregions and river basins linked to domestic consumption and exports to China, EU, France, and the rest of the world, before quantifying damage to biodiversity, and water scarcity footprints. The EU had the lowest potential biodiversity damage but the largest water scarcity footprint following respective sourcing patterns in 12 ecoregions and 18 river basins. These results differed from the average impact scores obtained from Brazilian soybean production information alone. The CSM can be derived at different scales (subnationally, internationally) using existing supply chain information and constitutes an additional step toward greater regionalization in LCAs, particularly for impacts with greater spatial variability such as biodiversity and water scarcity.

Second rate or a second chance? Assessing biomass and biodiversity recovery in regenerating Amazonian forests.

Secondary forests (SFs) regenerating on previously deforested land account for large, expanding areas of tropical forest cover. Given that tropical forests rank among Earth's most important reservoirs of carbon and biodiversity, SFs play an increasingly pivotal role in the carbon cycle and as potential habitat for forest biota. Nevertheless, their capacity to regain the biotic attributes of undisturbed primary forests (UPFs) remains poorly understood. Here, we provide a comprehensive assessment of SF recovery, using extensive tropical biodiversity, biomass, and environmental datasets. These data, collected in 59 naturally regenerating SFs and 30 co-located UPFs in the eastern Amazon, cover >1,600 large- and small-stemmed plant, bird, and dung beetles species and a suite of forest structure, landscape context, and topoedaphic predictors. After up to 40 years of regeneration, the SFs we surveyed showed a high degree of biodiversity resilience, recovering, on average among taxa, 88% and 85% mean UPF species richness and composition, respectively. Across the first 20 years of succession, the period for which we have accurate SF age data, biomass recovered at 1.2% per year, equivalent to a carbon uptake rate of 2.25 Mg/ha per year, while, on average, species richness and composition recovered at 2.6% and 2.3% per year, respectively. For all taxonomic groups, biomass was strongly associated with SF species distributions. However, other variables describing habitat complexity-canopy cover and understory stem density-were equally important occurrence predictors for most taxa. Species responses to biomass revealed a successional transition at approximately 75 Mg/ha, marking the influx of high-conservation-value forest species. Overall, our results show that naturally regenerating SFs can accumulate substantial amounts of carbon and support many forest species. However, given that the surveyed SFs failed to return to a typical UPF state, SFs are not substitutes for UPFs.

Disentangling the pathways of land use impacts on the functional structure of fish assemblages in Amazon streams.

Agricultural land use is a primary driver of environmental impacts on streams. However, the causal processes that shape these impacts operate through multiple pathways and at several spatial scales. This complexity undermines the development of more effective management approaches, and illustrates the need for more in-depth studies to assess the mechanisms that determine changes in stream biodiversity. Here we present results of the most comprehensive multi-scale assessment of the biological condition of streams in the Amazon to date, examining functional responses of fish assemblages to land use. We sampled fish assemblages from two large human-modified regions, and characterized stream conditions by physical habitat attributes and key landscape-change variables, including density of road crossings (i.e. riverscape fragmentation), deforestation, and agricultural intensification. Fish species were functionally characterized using ecomorphological traits describing feeding, locomotion, and habitat preferences, and these traits were used to derive indices that quantitatively describe the functional structure of the assemblages. Using structural equation modeling, we disentangled multiple drivers operating at different spatial scales, identifying causal pathways that significantly affect stream condition and the structure of the fish assemblages. Deforestation at catchment and riparian network scales altered the channel morphology and the stream bottom structure, changing the functional identity of assemblages. Local deforestation reduced the functional evenness of assemblages (i.e. increased dominance of specific trait combinations) mediated by expansion of aquatic vegetation cover. Riverscape fragmentation reduced functional richness, evenness and divergence, suggesting a trend toward functional homogenization and a reduced range of ecological niches within assemblages following the loss of regional connectivity. These results underscore the often-unrecognized importance of different land use changes, each of which can have marked effects on stream biodiversity. We draw on the relationships observed herein to suggest priorities for the improved management of stream systems in the multiple-use landscapes that predominate in human-modified tropical forests.

Primary forests are irreplaceable for sustaining tropical biodiversity.

Human-driven land-use changes increasingly threaten biodiversity, particularly in tropical forests where both species diversity and human pressures on natural environments are high. The rapid conversion of tropical forests for agriculture, timber production and other uses has generated vast, human-dominated landscapes with potentially dire consequences for tropical biodiversity. Today, few truly undisturbed tropical forests exist, whereas those degraded by repeated logging and fires, as well as secondary and plantation forests, are rapidly expanding. Here we provide a global assessment of the impact of disturbance and land conversion on biodiversity in tropical forests using a meta-analysis of 138 studies. We analysed 2,220 pairwise comparisons of biodiversity values in primary forests (with little or no human disturbance) and disturbed forests. We found that biodiversity values were substantially lower in degraded forests, but that this varied considerably by geographic region, taxonomic group, ecological metric and disturbance type. Even after partly accounting for confounding colonization and succession effects due to the composition of surrounding habitats, isolation and time since disturbance, we find that most forms of forest degradation have an overwhelmingly detrimental effect on tropical biodiversity. Our results clearly indicate that when it comes to maintaining tropical biodiversity, there is no substitute for primary forests.

Actor-specific contributions to the deforestation slowdown in the Brazilian Amazon.

Annual deforestation rates in the Brazilian Amazon fell by 77% between 2004 and 2011, yet have stabilized since 2009 at 5,000-7,000 km(2). We provide the first submunicipality assessment, to our knowledge, of actor-specific contributions to the deforestation slowdown by linking agricultural census and remote-sensing data on deforestation and forest degradation. Almost half (36,158 km(2)) of the deforestation between 2004 and 2011 occurred in areas dominated by larger properties (>500 ha), whereas only 12% (9,720 km(2)) occurred in areas dominated by smallholder properties (<100 ha). In addition, forests in areas dominated by smallholders tend to be less fragmented and less degraded. However, although annual deforestation rates fell during this period by 68-85% for all actors, the contribution of the largest landholders (>2,500 ha) to annual deforestation decreased over time (63% decrease between 2005 and 2011), whereas that of smallholders went up by a similar amount (69%) during the same period. In addition, the deforestation share attributable to remote areas increased by 88% between 2009 and 2011. These observations are consistent across the Brazilian Amazon, regardless of geographical differences in actor dominance or socioenvironmental context. Our findings suggest that deforestation policies to date, which have been particularly focused on command and control measures on larger properties in deforestation hotspots, may be increasingly limited in their effectiveness and fail to address all actors equally. Further reductions in deforestation are likely to be increasingly costly and require actor-tailored approaches, including better monitoring to detect small-scale deforestation and a shift toward more incentive-based conservation policies.

Land use intensification in the humid tropics increased both alpha and beta diversity of soil bacteria.

Anthropogenic pressures on tropical forests are rapidly intensifying, but our understanding of their implications for biological diversity is still very limited, especially with regard to soil biota, and in particular soil bacterial communities. Here we evaluated bacterial community composition and diversity across a gradient of land use intensity in the eastern Amazon from undisturbed primary forest, through primary forests varyingly disturbed by fire, regenerating secondary forest, pasture, and mechanized agriculture. Soil bacteria were assessed by paired-end Illumina sequencing of 16S rRNA gene fragments (V4 region). The resulting sequences were clustered into operational taxonomic units (OTU) at a 97% similarity threshold. Land use intensification increased the observed bacterial diversity (both OTU richness and community heterogeneity across space) and this effect was strongly associated with changes in soil pH. Moreover, land use intensification and subsequent changes in soil fertility, especially pH, altered the bacterial community composition, with pastures and areas of mechanized agriculture displaying the most contrasting communities in relation to undisturbed primary forest. Together, these results indicate that tropical forest conversion impacts soil bacteria not through loss of diversity, as previously thought, but mainly by imposing marked shifts on bacterial community composition, with unknown yet potentially important implications for ecological functions and services performed by these communities.

Idiosyncratic responses of Amazonian birds to primary forest disturbance.

As humans continue to alter tropical landscapes across the world, it is important to understand what environmental factors help determine the persistence of biodiversity in modified ecosystems. Studies on well-known taxonomic groups can offer critical insights as to the fate of biodiversity in these modified systems. Here we investigated species-specific responses of 44 forest-associated bird species with different behavioural traits to forest disturbance in 171 transects distributed across 31 landscapes in two regions of the eastern Brazilian Amazon. We investigated patterns of species occurrence in primary forests varyingly disturbed by selective-logging and fire and examined the relative importance of local, landscape and historical environmental variables in determining species occurrences. Within undisturbed and disturbed primary forest transects, we found that distance to forest edge and the biomass of large trees were the most important predictors driving the occurrence of individual species. However, we also found considerable variation in species responses to different environmental variables as well as inter-regional variation in the responses of the same species to the same environmental variables. We advocate the utility of using species-level analyses to complement community-wide responses in order to uncover highly variable and species-specific responses to environmental change that remain so poorly understood.

How pervasive is biotic homogenization in human-modified tropical forest landscapes?

Land-cover change and ecosystem degradation may lead to biotic homogenization, yet our understanding of this phenomenon over large spatial scales and different biotic groups remains weak. We used a multi-taxa dataset from 335 sites and 36 heterogeneous landscapes in the Brazilian Amazon to examine the potential for landscape-scale processes to modulate the cumulative effects of local disturbances. Biotic homogenization was high in production areas but much less in disturbed and regenerating forests, where high levels of among-site and among-landscape ß-diversity appeared to attenuate species loss at larger scales. We found consistently high levels of ß-diversity among landscapes for all land cover classes, providing support for landscape-scale divergence in species composition. Our findings support concerns that ß-diversity has been underestimated as a driver of biodiversity change and underscore the importance of maintaining a distributed network of reserves, including remaining areas of undisturbed primary forest, but also disturbed and regenerating forests, to conserve regional biota.

Thresholds of species loss in Amazonian deforestation frontier landscapes.

In the Brazilian Amazon, private land accounts for the majority of remaining native vegetation. Understanding how land-use change affects the composition and distribution of biodiversity in farmlands is critical for improving conservation strategies in the face of rapid agricultural expansion. Working across an area exceeding 3 million ha in the southwestern state of Rondônia, we assessed how the extent and configuration of remnant forest in replicate 10,000-ha landscapes has affected the occurrence of a suite of Amazonian mammals and birds. In each of 31 landscapes, we used field sampling and semistructured interviews with landowners to determine the presence of 28 large and medium sized mammals and birds, as well as a further 7 understory birds. We then combined results of field surveys and interviews with a probabilistic model of deforestation. We found strong evidence for a threshold response of sampled biodiversity to landscape level forest cover; landscapes with <30-40% forest cover hosted markedly fewer species. Results from field surveys and interviews yielded similar thresholds. These results imply that in partially deforested landscapes many species are susceptible to extirpation following relatively small additional reductions in forest area. In the model of deforestation by 2030 the number of 10,000-ha landscapes under a conservative threshold of 43% forest cover almost doubled, such that only 22% of landscapes would likely to be able to sustain at least 75% of the 35 focal species we sampled. Brazilian law requires rural property owners in the Amazon to retain 80% forest cover, although this is rarely achieved. Prioritizing efforts to ensure that entire landscapes, rather than individual farms, retain at least 50% forest cover may help safeguard native biodiversity in private forest reserves in the Amazon.

Road networks predict human influence on Amazonian bird communities.

Road building can lead to significant deleterious impacts on biodiversity, varying from direct road-kill mortality and direct habitat loss associated with road construction, to more subtle indirect impacts from edge effects and fragmentation. However, little work has been done to evaluate the specific effects of road networks and biodiversity loss beyond the more generalized effects of habitat loss. Here, we compared forest bird species richness and composition in the municipalities of Santarém and Belterra in Pará state, eastern Brazilian Amazon, with a road network metric called 'roadless volume (RV)' at the scale of small hydrological catchments (averaging 3721 ha). We found a significant positive relationship between RV and both forest bird richness and the average number of unique species (species represented by a single record) recorded at each site. Forest bird community composition was also significantly affected by RV. Moreover, there was no significant correlation between RV and forest cover, suggesting that road networks may impact biodiversity independently of changes in forest cover. However, variance partitioning analysis indicated that RV has partially independent and therefore additive effects, suggesting that RV and forest cover are best used in a complementary manner to investigate changes in biodiversity. Road impacts on avian species richness and composition independent of habitat loss may result from road-dependent habitat disturbance and fragmentation effects that are not captured by total percentage habitat cover, such as selective logging, fire, hunting, traffic disturbance, edge effects and road-induced fragmentation.

Two hundred years of local avian extinctions in eastern Amazonia.

Local, regional, and global extinctions caused by habitat loss, degradation, and fragmentation have been widely reported for the tropics. The patterns and drivers of this loss of species are now increasingly well known in Amazonia, but there remains a significant gap in understanding of long-term trends in species persistence and extinction in anthropogenic landscapes. Such a historical perspective is critical for understanding the status and trends of extant biodiversity as well as for identifying priorities to halt further losses. Using extensive historical data sets of specimen records and results of contemporary surveys, we searched for evidence of local extinctions of a terra firma rainforest avifauna over 200 years in a 2500 km(2) eastern Amazonian region around the Brazilian city of Belém. This region has the longest history of ornithological fieldwork in the entire Amazon basin and lies in the highly threatened Belém Centre of Endemism. We also compared our historically inferred extinction events with extensive data on species occurrences in a sample of catchments in a nearby municipality (Paragominas) that encompass a gradient of past forest loss. We found evidence for the possible extinction of 47 species (14% of the regional species pool) that were unreported from 1980 to 2013 (80% last recorded between 1900 and 1980). Seventeen species appear on the International Union for Conservation of Nature Red List, and many of these are large-bodied. The species lost from the region immediately around Belém are similar to those which are currently restricted to well-forested catchments in Paragominas. Although we anticipate the future rediscovery or recolonization of some species inferred to be extinct by our calculations, we also expect that there are likely to be additional local extinctions, not reported here, given the ongoing loss and degradation of remaining areas of native vegetation across eastern Amazonia.

Biodiversity offsets and the challenge of achieving no net loss.

Businesses, governments, and financial institutions are increasingly adopting a policy of no net loss of biodiversity for development activities. The goal of no net loss is intended to help relieve tension between conservation and development by enabling economic gains to be achieved without concomitant biodiversity losses. biodiversity offsets represent a necessary component of a much broader mitigation strategy for achieving no net loss following prior application of avoidance, minimization, and remediation measures. However, doubts have been raised about the appropriate use of biodiversity offsets. We examined what no net loss means as a desirable conservation outcome and reviewed the conditions that determine whether, and under what circumstances, biodiversity offsets can help achieve such a goal. We propose a conceptual framework to substitute the often ad hoc approaches evident in many biodiversity offset initiatives. The relevance of biodiversity offsets to no net loss rests on 2 fundamental premises. First, offsets are rarely adequate for achieving no net loss of biodiversity alone. Second, some development effects may be too difficult or risky, or even impossible, to offset. To help to deliver no net loss through biodiversity offsets, biodiversity gains must be comparable to losses, be in addition to conservation gains that may have occurred in absence of the offset, and be lasting and protected from risk of failure. Adherence to these conditions requires consideration of the wider landscape context of development and offset activities, timing of offset delivery, measurement of biodiversity, accounting procedures and rule sets used to calculate biodiversity losses and gains and guide offset design, and approaches to managing risk. Adoption of this framework will strengthen the potential for offsets to provide an ecologically defensible mechanism that can help reconcile conservation and development. Balances de Biodiversidad y el Reto de No Obtener Pérdida Neta.

Disentangling the numbers behind agriculture-driven tropical deforestation.

Tropical deforestation continues at alarming rates with profound impacts on ecosystems, climate, and livelihoods, prompting renewed commitments to halt its continuation. Although it is well established that agriculture is a dominant driver of deforestation, rates and mechanisms remain disputed and often lack a clear evidence base. We synthesize the best available pantropical evidence to provide clarity on how agriculture drives deforestation. Although most (90 to 99%) deforestation across the tropics 2011 to 2015 was driven by agriculture, only 45 to 65% of deforested land became productive agriculture within a few years. Therefore, ending deforestation likely requires combining measures to create deforestation-free supply chains with landscape governance interventions. We highlight key remaining evidence gaps including deforestation trends, commodity-specific land-use dynamics, and data from tropical dry forests and forests across Africa.

The timing and causality of ecological shifts on Caribbean reefs.

Caribbean reefs have experienced unprecedented changes in the past four decades. Of great concern is the perceived widespread shift from coral to macroalgal dominance and the question of whether it represents a new, stable equilibrium for coral-reef communities. The primary causes of the shift-grazing pressure (top-down), nutrient loading (bottom-up) or direct coral mortality (side-in)-still remain somewhat controversial in the coral-reef literature. We have attempted to tease out the relative importance of each of these causes. Four insights emerge from our analysis of an early regional dataset of information on the benthic composition of Caribbean reefs spanning the years 1977-2001. First, although three-quarters of reef sites have experienced coral declines concomitant with macroalgal increases, fewer than 10% of the more than 200 sites studied were dominated by macroalgae in 2001, by even the most conservative definition of dominance. Using relative dominance as the threshold, a total of 49 coral-to-macroalgae shifts were detected. This total represents ~35% of all sites that were dominated by coral at the start of their monitoring periods. Four shifts (8.2%) occurred because of coral loss with no change in macroalgal cover, 15 (30.6%) occurred because of macroalgal gain without coral loss, and 30 (61.2%) occurred owing to concomitant coral decline and macroalgal increase. Second, the timing of shifts at the regional scale is most consistent with the side-in model of reef degradation, which invokes coral mortality as a precursor to macroalgal takeover, because more shifts occurred after regional coral-mortality events than expected by chance. Third, instantaneous observations taken at the start and end of the time-series for individual sites showed these reefs existed along a continuum of coral and macroalgal cover. The continuous, broadly negative relationship between coral and macroalgal cover suggests that in some cases coral-to-macroalgae phase shifts may be reversed by removing sources of perturbation or restoring critical components such as the herbivorous sea urchin Diadema antillarum to the system. The five instances in which macroalgal dominance was reversed corroborate the conclusion that macroalgal dominance is not a stable, alternative community state as has been commonly assumed. Fourth, the fact that the loss in regional coral cover and concomitant changes to the benthic community are related to punctuated, discrete events with known causes (i.e. coral disease and bleaching), lends credence to the hypothesis that coral reefs of the Caribbean have been under assault from climate-change-related maladies since the 1970s.

Author Correction: Strategic approaches to restoring ecosystems can triple conservation gains and halve costs.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Integrated terrestrial-freshwater planning doubles conservation of tropical aquatic species.

Conservation initiatives overwhelmingly focus on terrestrial biodiversity, and little is known about the freshwater cobenefits of terrestrial conservation actions. We sampled more than 1500 terrestrial and freshwater species in the Amazon and simulated conservation for species from both realms. Prioritizations based on terrestrial species yielded on average just 22% of the freshwater benefits achieved through freshwater-focused conservation. However, by using integrated cross-realm planning, freshwater benefits could be increased by up to 600% for a 1% reduction in terrestrial benefits. Where freshwater biodiversity data are unavailable but aquatic connectivity is accounted for, freshwater benefits could still be doubled for negligible losses of terrestrial coverage. Conservation actions are urgently needed to improve the status of freshwater species globally. Our results suggest that such gains can be achieved without compromising terrestrial conservation goals.

Prospects for tropical forest biodiversity in a human-modified world.

The future of tropical forest biodiversity depends more than ever on the effective management of human-modified landscapes, presenting a daunting challenge to conservation practitioners and land use managers. We provide a critical synthesis of the scientific insights that guide our understanding of patterns and processes underpinning forest biodiversity in the human-modified tropics, and present a conceptual framework that integrates a broad range of social and ecological factors that define and contextualize the possible future of tropical forest species. A growing body of research demonstrates that spatial and temporal patterns of biodiversity are the dynamic product of interacting historical and contemporary human and ecological processes. These processes vary radically in their relative importance within and among regions, and have effects that may take years to become fully manifest. Interpreting biodiversity research findings is frequently made difficult by constrained study designs, low congruence in species responses to disturbance, shifting baselines and an over-dependence on comparative inferences from a small number of well studied localities. Spatial and temporal heterogeneity in the potential prospects for biodiversity conservation can be explained by regional differences in biotic vulnerability and anthropogenic legacies, an ever-tighter coupling of human-ecological systems and the influence of global environmental change. These differences provide both challenges and opportunities for biodiversity conservation. Building upon our synthesis we outline a simple adaptive-landscape planning framework that can help guide a new research agenda to enhance biodiversity conservation prospects in the human-modified tropics.