Barry W. Brook

Synergies among extinction drivers under global change

If habitat destruction or overexploitation of populations is severe, species loss can occur directly and abruptly. Yet the final descent to extinction is often driven by synergistic processes (amplifying feedbacks) that can be disconnected from the original cause of decline. We review recent observational, experimental and meta-analytic work which together show that owing to interacting and self-reinforcing processes, estimates of extinction risk for most species are more severe than previously recognised. As such, conservation actions which only target single-threat drivers risk being inadequate because of the cascading effects caused by unmanaged synergies. Future work should focus on how climate change will interact with and accelerate ongoing threats to biodiversity, such as habitat degradation, overexploitation and invasive species.

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.

Catastrophic extinctions follow deforestation in Singapore

The looming mass extinction of biodiversity in the humid tropics is a major concern for the future, yet most reports of extinctions in these regions are anecdotal or conjectural, with a scarcity of robust, broad-based empirical data. Here we report on local extinctions among a wide range of terrestrial and freshwater taxa from Singapore (540 km2) in relation to habitat loss exceeding 95% over 183 years. Substantial rates of documented and inferred extinctions were found, especially for forest specialists, with the greatest proportion of extinct taxa (34–87%) in butterflies, fish, birds and mammals. Observed extinctions were generally fewer, but inferred losses often higher, in vascular plants, phasmids, decapods, amphibians and reptiles (5–80%). Forest reserves comprising only 0.25% of Singapores area now harbour over 50% of the residual native biodiversity. Extrapolations of the observed and inferred local extinction data, using a calibrated species–area model, imply that the current unprecedented rate of habitat destruction in Southeast Asia will result in the loss of 13–42% of regional populations over the next century, at least half of which will represent global species extinctions.

PREDICTIVE ACCURACY OF POPULATION VIABILITY ANALYSIS IN CONSERVATION BIOLOGY

Population viability analysis (PVA) is widely applied in conservation biology to predict extinction risks for threatened species and to compare alternative options for their mangement. It can also be used as a basis for listing species as endangered under World Conservation Union criteria. However, there is considerable scepticism regarding the predictive accuracy of PVA, mainly because of a lack of validation in real systems. Here we conducted a retrospective test of PVA based on 21 long-term ecological studies—the first comprehensive and replicated evaluation of the predictive powers of PVA. Parameters were estimated from the first half of each data set and the second half was used to evaluate the performance of the model. Contrary to recent criticisms, we found that PVA predictions were surprisingly accurate. The risk of population decline closely matched observed outcomes, there was no significant bias, and population size projections did not differ significantly from reality. Furthermore, the predictions of the five PVA software packages were highly concordant. We conclude that PVA is a valid and sufficiently accurate tool for categorizing and managing endangered species.

Estimates of minimum viable population sizes for vertebrates and factors influencing those estimates

Population size is a major determinant of extinction risk. However, controversy remains as to how large populations need to be to ensure persistence. It is generally believed that minimum viable population sizes (MVPs) would be highly specific, depending on the environmental and life history characteristics of the species. We used population viability analysis to estimate MVPs for 102 species. We define a minimum viable population size as one with a 99% probability of persistence for 40 generations. The models are comprehensive and include age-structure, catastrophes, demographic stochasticity, environmental stochasticity, and inbreeding depression. The mean and median estimates of MVP were 7316 and 5816 adults, respectively. This is slightly larger than, but in general agreement with, previous estimates of MVP. MVPs did not differ significantly among major taxa, or with latitude or trophic level, but were negatively correlated with population growth rate and positively correlated with the length of the study used to parameterize the model. A doubling of study duration increased the estimated MVP by approximately 67%. The increase in extinction risk is associated with greater temporal variation in population size for models built from longer data sets. Short-term studies consistently underestimate the true variances for demographic parameters in populations. Thus, the lack of long-term studies for endangered species leads to widespread underestimation of extinction risk. The results of our simulations suggest that conservation programs, for wild populations, need to be designed to conserve habitat capable of supporting approximately 7000 adult vertebrates in order to ensure long-term persistence. # 2003 Elsevier Science Ltd. All rights reserved.

Measuring the Meltdown: Drivers of Global Amphibian Extinction and Decline

Habitat loss, climate change, over-exploitation, disease and other factors have been hypothesised in the global decline of amphibian biodiversity. However, the relative importance of and synergies among different drivers are still poorly understood. We present the largest global analysis of roughly 45% of known amphibians (2,583 species) to quantify the influences of life history, climate, human density and habitat loss on declines and extinction risk. Multi-model Bayesian inference reveals that large amphibian species with small geographic range and pronounced seasonality in temperature and precipitation are most likely to be Red-Listed by IUCN. Elevated habitat loss and human densities are also correlated with high threat risk. Range size, habitat loss and more extreme seasonality in precipitation contributed to decline risk in the 2,454 species that declined between 1980 and 2004, compared to species that were stable (n = 1,545) or had increased (n = 28). These empirical results show that amphibian species with restricted ranges should be urgently targeted for conservation.

The state and conservation of Southeast Asian biodiversity.

Southeast Asia is a region of conservation concern due to heavy losses of its native habitats. In this overview, we highlight the conservation importance of Southeast Asia by comparing its degree of species endemism and endangerment, and its rate of deforestation with other tropical regions (i.e., Meso-America, South America, and Sub-Saharan Africa). Southeast Asia contains the highest mean proportion of country-endemic bird (9%) and mammal species (11%). This region also has the highest proportion of threatened vascular plant, reptile, bird, and mammal species. Furthermore, not only is Southeast Asia’s annual deforestation rate the highest in the tropics, but it has also increased between the periods 1990–2000 and 2000–2005. This could result in projected losses of 13–85% of biodiversity in the region by 2100. Secondary habitat restoration, at least in certain countries, would allow for some amelioration of biodiversity loss and thus potentially lower the currently predicted extinction rates. Nonetheless, urgent conservation actions are needed. Conservation initiatives should include public education, sustaining livelihoods, and ways to enhance the sustainability of agriculture and increase the capacity of conservation institutions. Furthermore, these actions should be country-specific and not ignore areas heavily populated by humans, as they can also harbour high numbers of threatened species. We urge that cooperative conservation initiatives be undertaken and support (e.g., capacity-building) be given by more developed countries in the region and beyond.

STRENGTH OF EVIDENCE FOR DENSITY DEPENDENCE IN ABUNDANCE TIME SERIES OF 1198 SPECIES

Population limitation is a fundamental tenet of ecology, but the relative roles of exogenous and endogenous mechanisms remain unquantified for most species. Here we used multi-model inference (MMI), a form of model averaging, based on information theory (Akaikes Information Criterion) to evaluate the relative strength of evidence for density-dependent and density-independent population dynamical models in long-term abundance time series of 1198 species. We also compared the MMI results to more classic methods for detecting density dependence: Neyman-Pearson hypothesis-testing and best-model selection using the Bayesian Information Criterion or cross-validation. Using MMI on our large database, we show that density dependence is a pervasive feature of population dynamics (median MMI support for density dependence = 74.7-92.2%), and that this holds across widely different taxa. The weight of evidence for density dependence varied among species but increased consistently, with the number of generations monitored. Best-model selection methods yielded similar results to MMI (a density-dependent model was favored in 66.2-93.9% of species time series), while the hypothesis-testing methods detected density dependence less frequently (32.6-49.8%). There were no obvious differences in the prevalence of density dependence across major taxonomic groups under any of the statistical methods used. These results underscore the value of using multiple modes of analysis to quantify the relative empirical support for a set of working hypotheses that encompass a range of realistic population dynamical behaviors.

Tropical turmoil: a biodiversity tragedy in progress

All is not well for biodiversity in the tropics. Despite recent debate over the extent of future tropical extinctions and the effectiveness of reserve systems, the continued disappearance of habitat, soaring human population, and loss of vital ecosystem services demand immediate action. This crisis is worrying, given that tropical regions support over two-thirds of all known species and are populated by some of the worlds poorest people, who have little recourse to lower environmental-impact lifestyles. Recent evidence has shown that – in addition to unabated rates of forest loss – coastal development, overexploitation of wildlife, catchment modification, and habitat conversion are threatening human well-being. We argue that the recent technical debate about likely extinctions masks the real issue – that, to prevent further loss of irreplaceable tropical biodiversity, we must err on the side of caution. We need to avoid inadvertently supporting political agendas that assume low future extinction rates, because this will result in further destruction of tropical biodiversity.

Contribution of inbreeding to extinction risk in threatened species

Wild populations face threats both from deterministic factors, e.g., habitat loss, overexploitation, pollution, and introduced species, and from stochastic events of a demographic, genetic, and environmental nature, including catastrophes. Inbreeding reduces reproductive fitness in naturally outbreeding species, but its role in extinctions of wild populations is controversial. To evaluate critically the role of inbreeding in extinction, we conducted realistic population viability analyses of 20 threatened species, with and without inbreeding depression, using initial population sizes of 50, 250, and 1000. Inbreeding markedly decreased median times to extinction by 28.5, 30.5, and 25% for initial populations of 50, 250, and 1000, respectively, and the impacts were similar across major taxa. The major variable explaining differences among species was initial population growth rate, whereas the impact of inbreeding was least in species with negative growth rates. These results demonstrate that the prospects for survival of threatened species will usually be seriously overestimated if genetic factors are disregarded, and that inappropriate recovery plans may be instituted if inbreeding depression is ignored.