Global economic trade-offs between wild nature and tropical agriculture.

Global demands for agricultural and forestry products provide economic incentives for deforestation across the tropics. Much of this deforestation occurs with a lack of information on the spatial distribution of benefits and costs of deforestation. To inform global sustainable land-use policies, we combine geographic information systems (GIS) with a meta-analysis of ecosystem services (ES) studies to perform a spatially explicit analysis of the trade-offs between agricultural benefits, carbon emissions, and losses of multiple ecosystem services because of tropical deforestation from 2000 to 2012. Even though the value of ecosystem services presents large inherent uncertainties, we find a pattern supporting the argument that the externalities of destroying tropical forests are greater than the current direct economic benefits derived from agriculture in all cases bar one: when yield and rent potentials of high-value crops could be realized in the future. Our analysis identifies the Atlantic Forest, areas around the Gulf of Guinea, and Thailand as areas where agricultural conversion appears economically efficient, indicating a major impediment to the long-term financial sustainability of Reducing Emissions from Deforestation and forest Degradation (REDD+) schemes in those countries. By contrast, Latin America, insular Southeast Asia, and Madagascar present areas with low agricultural rents (ARs) and high values in carbon stocks and ES, suggesting that they are economically viable conservation targets. Our study helps identify optimal areas for conservation and agriculture together with their associated uncertainties, which could enhance the efficiency and sustainability of pantropical land-use policies and help direct future research efforts.

Increasing arboreality with altitude: a novel biogeographic dimension.

Biodiversity is spatially organized by climatic gradients across elevation and latitude. But do other gradients exist that might drive biogeographic patterns? Here, we show that rainforest's vertical strata provide climatic gradients much steeper than those offered by elevation and latitude, and biodiversity of arboreal species is organized along this gradient. In Philippine and Singaporean rainforests, we demonstrate that rainforest frogs tend to shift up in the rainforest strata as altitude increases. Moreover, a Philippine-wide dataset of frog distributions shows that frog assemblages become increasingly arboreal at higher elevations. Thus, increased arboreality with elevation at broad biogeographic scales mirrors patterns we observed at local scales. Our proposed 'arboreality hypothesis' suggests that the ability to exploit arboreal habitats confers the potential for larger geographical distributions because species can shift their location in the rainforest strata to compensate for shifts in temperature associated with elevation and latitude. This novel finding may help explain patterns of species richness and abundance wherever vegetation produces a vertical microclimatic gradient. Our results further suggest that global warming will 'flatten' the biodiversity in rainforests by pushing arboreal species towards the cooler and wetter ground. This 'flattening' could potentially have serious impacts on forest functioning and species survival.

Reservoirs of richness: least disturbed tropical forests are centres of undescribed species diversity.

In the last few decades, there has been a remarkable discovery of new species of plants, invertebrates and vertebrates, in what have been called the new age of discovery. However, owing to anthropogenic impacts such as habitat conversion, many of the still unknown species may go extinct before being scientifically documented (i.e. 'crypto-extinctions'). Here, by applying a mathematical model of species descriptions which accounts for taxonomic effort, we show that even after 250 years of taxonomic classification, about 3050 amphibians and at least 160 land mammal species remain to be discovered and described. These values represent, respectively, 33 and 3 per cent of the current species total for amphibians and land mammals. We found that tropical moist forests of the Neotropics, Afrotropics and Indomalaya probably harbour the greatest numbers of undescribed species. Tropical forests with minimal anthropogenic disturbance are predicted to have larger proportions of undescribed species. However, the protected area coverage is low in many of these key biomes. Moreover, undescribed species are likely to be at a greater risk of extinction compared with known species because of small geographical ranges among other factors. By highlighting the key areas of undescribed species diversity, our study provides a starting template to rapidly document these species and protect them through better habitat management.

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.

Conservation successes at micro-, meso- and macroscales.

Although large-scale biodiversity declines are ongoing, certain conservation actions have made a positive difference. Rates of extinction and endangerment of vertebrate species, for instance, have probably been reduced via conservation interventions. Such conservation actions operate at different spatial scales. Habitat preservation and endangered species recovery are examples of conservation successes at microscales. Mesoscale conservation includes regional cooperation among neighboring countries that has arrested population declines of endangered species, such as mountain gorillas. At macroscales, public pressure on multinational corporations has sometimes resulted in their abandoning environmentally damaging practices or suppliers with poor environmental records. Overall, conservation projects such as these need more long-term funding and greater political and popular support, and must also include provisions to evaluate and document their outcomes. As we discuss here, a focus on conservation successes achieved at different scales can help to promote these aims and guide future conservation victories.

The world's rediscovered species: back from the brink?

Each year, numerous species thought to have disappeared are rediscovered. Yet, do these rediscoveries represent the return of viable populations or the delayed extinction of doomed species? We document the number, distribution and conservation status of rediscovered amphibian, bird, and mammal species globally. Over the past 122 years, at least 351 species have been rediscovered, most occurring in the tropics. These species, on average, were missing for 61 years before being rediscovered (range of 3-331 years). The number of rediscoveries per year increased over time and the majority of these rediscoveries represent first documentations since their original description. Most rediscovered species have restricted ranges and small populations, and 92% of amphibians, 86% of birds, and 86% of mammals are highly threatened, independent of how long they were missing or when they were rediscovered. Under the current trends of widespread habitat loss, particularly in the tropics, most rediscovered species remain on the brink of extinction.

Endemic predators, invasive prey and native diversity.

Interactions between native diversity and invasive species can be more complex than is currently understood. Invasive ant species often substantially reduce diversity in the native ants diversity that act as natural control agents for pest insects. In Indonesia (on the island of Sulawesi), the third largest cacao producer worldwide, we show that a predatory endemic toad (Ingerophrynus celebensis) controls invasive ant (Anoplolepis gracilipes) abundance, and positively affects native ant diversity. We call this the invasive-naivety effect (an opposite of enemy release), whereby alien species may not harbour anti-predatory defences against a novel native predator. A positive effect of the toads on native ants may facilitate their predation on insect vectors of cacao diseases. Hence, toads may increase crop yield, but further research is needed on this aspect. Ironically, amphibians are globally the most threatened vertebrate class and are strongly impacted by the conversion of rainforest to cacao plantations in Sulawesi. It is, therefore, crucial to manage cacao plantations to maintain these endemic toads, as they may provide critical ecosystem services, such as invasion resistance and preservation of native insect diversity.

The human dimension of fire regimes on Earth.

Humans and their ancestors are unique in being a fire-making species, but 'natural' (i.e. independent of humans) fires have an ancient, geological history on Earth. Natural fires have influenced biological evolution and global biogeochemical cycles, making fire integral to the functioning of some biomes. Globally, debate rages about the impact on ecosystems of prehistoric human-set fires, with views ranging from catastrophic to negligible. Understanding of the diversity of human fire regimes on Earth in the past, present and future remains rudimentary. It remains uncertain how humans have caused a departure from 'natural' background levels that vary with climate change. Available evidence shows that modern humans can increase or decrease background levels of natural fire activity by clearing forests, promoting grazing, dispersing plants, altering ignition patterns and actively suppressing fires, thereby causing substantial ecosystem changes and loss of biodiversity. Some of these contemporary fire regimes cause substantial economic disruptions owing to the destruction of infrastructure, degradation of ecosystem services, loss of life, and smoke-related health effects. These episodic disasters help frame negative public attitudes towards landscape fires, despite the need for burning to sustain some ecosystems. Greenhouse gas-induced warming and changes in the hydrological cycle may increase the occurrence of large, severe fires, with potentially significant feedbacks to the Earth system. Improved understanding of human fire regimes demands: (1) better data on past and current human influences on fire regimes to enable global comparative analyses, (2) a greater understanding of different cultural traditions of landscape burning and their positive and negative social, economic and ecological effects, and (3) more realistic representations of anthropogenic fire in global vegetation and climate change models. We provide an historical framework to promote understanding of the development and diversification of fire regimes, covering the pre-human period, human domestication of fire, and the subsequent transition from subsistence agriculture to industrial economies. All of these phases still occur on Earth, providing opportunities for comparative research.

Evaluating the relative environmental impact of countries.

Environmental protection is critical to maintain ecosystem services essential for human well-being. It is important to be able to rank countries by their environmental impact so that poor performers as well as policy 'models' can be identified. We provide novel metrics of country-specific environmental impact ranks - one proportional to total resource availability per country and an absolute (total) measure of impact - that explicitly avoid incorporating confounding human health or economic indicators. Our rankings are based on natural forest loss, habitat conversion, marine captures, fertilizer use, water pollution, carbon emissions and species threat, although many other variables were excluded due to a lack of country-specific data. Of 228 countries considered, 179 (proportional) and 171 (absolute) had sufficient data for correlations. The proportional index ranked Singapore, Korea, Qatar, Kuwait, Japan, Thailand, Bahrain, Malaysia, Philippines and Netherlands as having the highest proportional environmental impact, whereas Brazil, USA, China, Indonesia, Japan, Mexico, India, Russia, Australia and Peru had the highest absolute impact (i.e., total resource use, emissions and species threatened). Proportional and absolute environmental impact ranks were correlated, with mainly Asian countries having both high proportional and absolute impact. Despite weak concordance among the drivers of environmental impact, countries often perform poorly for different reasons. We found no evidence to support the environmental Kuznets curve hypothesis of a non-linear relationship between impact and per capita wealth, although there was a weak reduction in environmental impact as per capita wealth increases. Using structural equation models to account for cross-correlation, we found that increasing wealth was the most important driver of environmental impact. Our results show that the global community not only has to encourage better environmental performance in less-developed countries, especially those in Asia, there is also a requirement to focus on the development of environmentally friendly practices in wealthier countries.

Mechanisms driving change: altered species interactions and ecosystem function through global warming.

1. We review the mechanisms behind ecosystem functions, the processes that facilitate energy transfer along food webs, and the major processes that allow the cycling of carbon, oxygen and nitrogen, and use case studies to show how these have already been, and will continue to be, altered by global warming. 2. Increased temperatures will affect the interactions between heterotrophs and autotrophs (e.g. pollination and seed dispersal), and between heterotrophs (e.g. predators-prey, parasites/pathogens-hosts), with generally negative ramifications for important ecosystem services (functions that provide direct benefit to human society such as pollination) and potential for heightened species co-extinction rates. 3. Mitigation of likely impacts of warming will require, in particular, the maintenance of species diversity as insurance for the provision of basic ecosystem services. Key to this will be long-term monitoring and focused research that seek to maintain ecosystem resilience in the face of global warming. 4. We provide guidelines for pursuing research that quantifies the nexus between ecosystem function and global warming. These include documentation of key functional species groups within systems, and understanding the principal outcomes arising from direct and indirect effects of a rapidly warming environment. Localized and targeted research and monitoring, complemented with laboratory work, will determine outcomes for resilience and guide adaptive conservation responses and long-term planning.

Deforestation and avian extinction on tropical landbridge islands.

There are few empirical data, particularly collected simultaneously from multiple sites, on extinctions resulting from human-driven land-use change. Southeast Asia has the highest deforestation rate in the world, but the resulting losses of biological diversity remain poorly documented. Between November 2006 and March 2008, we conducted bird surveys on six landbridge islands in Malaysia and Indonesia. These islands were surveyed previously for birds in the early 1900 s, when they were extensively forested. Our bird inventories of the islands were nearly complete, as indicated by sampling saturation curves and nonparametric true richness estimators. From zero (Pulau Malawali and Pulau Mantanani) to 15 (Pulau Bintan) diurnal resident landbird species were apparently extirpated since the early 1900 s. Adding comparable but published extinction data from Singapore to our regression analyses, we found there were proportionally fewer forest bird extinctions in areas with greater remaining forest cover. Nevertheless, the statistical evidence to support this relationship was weak, owing to our unavoidably small sample size. Bird species that are restricted to the Indomalayan region, lay few eggs, are heavier, and occupy a narrower habitat breadth, were most vulnerable to extinction on Pulau Bintan. This was the only island where sufficient data existed to analyze the correlates of extinction. Forest preservation and restoration are needed on these islands to conserve the remaining forest avifauna. Our study of landbridge islands indicates that deforestation may increasingly threaten Southeast Asian biodiversity.

Effects of land-use change on community composition of tropical amphibians and reptiles in Sulawesi, Indonesia.

Little is known about the effects of anthropogenic land-use change on the amphibians and reptiles of the biodiverse tropical forests of Southeast Asia. We studied a land-use modification gradient stretching from primary forest, secondary forest, natural-shade cacao agroforest, planted-shade cacao agroforest to open areas in central Sulawesi, Indonesia. We determined species richness, abundance, turnover, and community composition in all habitat types and related these to environmental correlates, such as canopy heterogeneity and thickness of leaf litter. Amphibian species richness decreased systematically along the land-use modification gradient, but reptile richness and abundance peaked in natural-shade cacao agroforests. Species richness and abundance patterns across the disturbance gradient were best explained by canopy cover and leaf-litter thickness in amphibians and by canopy heterogeneity and cover in reptiles. Amphibians were more severely affected by forest disturbance in Sulawesi than reptiles. Heterogeneous canopy cover and thick leaf litter should be maintained in cacao plantations to facilitate the conservation value for both groups. For long-term and sustainable use of plantations, pruned shade trees should be permanently kept to allow rejuvenation of cacao and, thus, to prevent repeated forest encroachment.

Urgent preservation of boreal carbon stocks and biodiversity.

Containing approximately one-third of all remaining global forests, the boreal ecosystem is a crucial store of carbon and a haven for diverse biological communities. Historically, fire and insects primarily drove the natural dynamics of this biome. However, human-mediated disturbances have increased in these forests during recent years, resulting in extensive forest loss for some regions, whereas others face heavy forest fragmentation or threat of exploitation. Current management practices are not likely to maintain the attendant boreal forest communities, nor are they adequate to mitigate climate change effects. There is an urgent need to preserve existing boreal forests and restore degraded areas if we are to avoid losing this relatively intact biodiversity haven and major global carbon sink.

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.

The sixth mass coextinction: are most endangered species parasites and mutualists?

The effects of species declines and extinction on biotic interactions remain poorly understood. The loss of a species is expected to result in the loss of other species that depend on it (coextinction), leading to cascading effects across trophic levels. Such effects are likely to be most severe in mutualistic and parasitic interactions. Indeed, models suggest that coextinction may be the most common form of biodiversity loss. Paradoxically, few historical or contemporary coextinction events have actually been recorded. We review the current knowledge of coextinction by: (i) considering plausible explanations for the discrepancy between predicted and observed coextinction rates; (ii) exploring the potential consequences of coextinctions; (iii) discussing the interactions and synergies between coextinction and other drivers of species loss, particularly climate change; and (iv) suggesting the way forward for understanding the phenomenon of coextinction, which may well be the most insidious threat to global biodiversity.