Trait‐based prediction of extinction risk of small‐bodied freshwater fishes

Small body size is generally correlated with r‐selected life‐history traits, including early maturation, short‐generation times, and rapid growth rates, that result in high population turnover and a reduced risk of extinction. Unlike other classes of vertebrates, however, small freshwater fishes appear to have an equal or greater risk of extinction than large fishes. We explored whether particular traits explain the International Union for Conservation of Nature (IUCN) Red List conservation status of small‐bodied freshwater fishes from 4 temperate river basins: Murray‐Darling, Australia; Danube, Europe; Mississippi‐Missouri, North America; and the Rio Grande, North America. Twenty‐three ecological and life‐history traits were collated for all 171 freshwater fishes of ≤120 mm total length. We used generalized linear mixed‐effects models to assess which combination of the 23 traits best explained whether a species was threatened or not threatened. We used the best models to predict the probability of 29 unclassified species being listed as threatened. With and without controlling for phylogeny at the family level, small body size—among small‐bodied species—was the most influential trait correlated with threatened species listings. The k‐folds cross‐validation demonstrated that body size and a random effect structure that included family predicted the threat status with an accuracy of 78% (SE 0.5). We identified 10 species likely to be threatened that are not listed as such on the IUCN Red List. Small body size is not a trait that provides universal resistance to extinction, particularly for vertebrates inhabiting environments affected by extreme habitat loss and fragmentation. We hypothesize that this is because small‐bodied species have smaller home ranges, lower dispersal capabilities, and heightened ecological specialization relative to larger vertebrates. Trait data and further model development are needed to predict the IUCN conservation status of the over 11,000 unclassified freshwater fishes, especially those under threat from proposed dam construction in the world's most biodiverse river basins.     

An Index of Risk as a Measure of Biodiversity Conservation Achieved through Land Reform

Abstract:  We measured the net progress of land reform in achieving a national policy goal for biodiversity conservation in the context of ongoing clearing of native vegetation and additions of land to a highly nonrepresentative (residual) reserve network, interior South Island, New Zealand. We used systematic conservation-planning approaches to develop a spatially explicit index of risk of biodiversity loss (RBL). The index incorporated information from national data sets that describe New Zealand's remaining indigenous land cover, legal protection, and land environments and modeled risk to biodiversity on the basis of stated assumptions about the effects of past habitat loss and legal protection. The index identified irreplaceable and vulnerable native habitats in lowland environments as the most at risk of biodiversity loss, and risk was correlated with the density of threatened plant records. To measure achievement, we used changes in the index that reflected gains made and opportunity costs incurred by legal protection and privatization. Application of the index to measure the difference made by land reform showed it had caused a net increase in the risk of biodiversity loss because most land vulnerable to habitat modification and rich in threatened plant species was privatized and land at least risk of biodiversity loss was protected. The application revealed that new high-elevation reserves did little to mitigate biodiversity decline, that privatization of low-elevation land further jeopardized the most vulnerable biodiversity in lowland native habitats, and that outcomes of land reform for biodiversity deteriorated over time. Further development of robust achievement measures is needed to encourage more accountable biodiversity conservation decisions.     

Effects of Future Infrastructure Development on Threat Status and Occurrence of Amazonian Birds

Abstract:  Researchers predict that new infrastructure development will sharply increase the rate and extent of deforestation in the Brazilian Amazon. There are no predictions, however, of which species it will affect. We used a spatially explicit model that predicts the location of deforestation in the Brazilian Amazon by 2020 on the basis of historical patterns of deforestation following infrastructure development. We overlaid the predicted deforested areas onto maps of bird ranges to estimate the amount of habitat loss within species ranges. We also estimated the amount of habitat loss within modified ecoregions, which were used as surrogates for areas of bird endemism. We then used the extent of occurrence criterion of the World Conservation Union to predict the future conservation status of birds in the Brazilian Amazon. At current rates of development, our results show that at least 16 species will qualify as threatened or will lose more than half of their forested habitat. We also identified several subspecies and isolated populations that would also qualify as threatened. Most of the taxa we identified are not currently listed as threatened, and the majority are associated with riverine habitats, which have been largely ignored in bird conservation in Amazonia. These habitats and the species they hold will be increasingly relevant to conservation as river courses are altered and hydroelectric dams are constructed in the Brazilian Amazon.     

A Comparative Analysis of Specialization and Extinction Risk in Temperate-Zone Bats

Abstract:  Identifying the factors that influence the extinction risk of animals is essential in conservation biology because they help identify endangered species and provide the basis for their preservation. We present a comparative study that uses data from the literature on the diet and morphological specialization of European and North American bat species to investigate the effect of specialization on extinction risk. We focused on bats because many species are endangered and their high ecological diversity makes them a good model system for our purpose. After correcting for phylogenetic inertia, we compared the influence of dietary niche breadth as a measure of food specialization and of wing morphology as a measure of foraging strategy, habitat adaptation, and migratory ability on the vulnerability of 35 insectivorous bat species. Our results do not support the hypothesis that a narrow dietary niche breadth is related to high extinction risk. Instead they suggest that habitat specialization, which is reflected in wing morphology, influences the extinction risk of bats. Our study shows that an initial risk assessment in temperate-zone bats could be based on data of wing morphology but not on dietary data obtained from fecal analyses.     

Determinants of bird conservation‐action implementation and associated population trends of threatened species

Conservation actions, such as habitat protection, attempt to halt the loss of threatened species and help their populations recover. The efficiency and the effectiveness of actions have been examined individually. However, conservation actions generally occur simultaneously, so the full suite of implemented conservation actions should be assessed. We used the conservation actions underway for all threatened and near‐threatened birds of the world (International Union for Conservation of Nature Red List of Threatened Species) to assess which biological (related to taxonomy and ecology) and anthropogenic (related to geoeconomics) factors were associated with the implementation of different classes of conservation actions. We also assessed which conservation actions were associated with population increases in the species targeted. Extinction‐risk category was the strongest single predictor of the type of conservation actions implemented, followed by landmass type (continent, oceanic island, etc.) and generation length. Species targeted by invasive nonnative species control or eradication programs, ex situ conservation, international legislation, reintroduction, or education, and awareness‐raising activities were more likely to have increasing populations. These results illustrate the importance of developing a predictive science of conservation actions and the relative benefits of each class of implemented conservation action for threatened and near‐threatened birds worldwide.     

Effects of roost specialization on extinction risk in bats

Understanding causes and consequences of ecological specialization is of major concern in conservation. Specialist species are particularly vulnerable to human activities. If their food or habitats are depleted or lost, they may not be able to exploit alternative resources, and population losses may result. We examined International Union for Conservation of Nature (IUCN) Red List bat data and the number of roosts used per species (accounting for phylogenetic independence) to determine whether roost specialization is correlated with extinction risk. We found a significant correlation between the IUCN Red List category and the number of roost types used. Species that use fewer roost types had a higher risk of extinction. We found that caves and similar structures were the most widely used roost types, particularly by species under some level of risk of extinction. Many critically endangered, endangered, or vulnerable species used natural roosts exclusively, whereas less threatened species used natural and human‐made roosts. Our results suggest that roost loss, particularly in species that rely on a single roost type, may be linked to extinction risk. Our focus on a single life history trait prevented us from determining how important this variable is for extinction risk relative to other variables, but we have taken a first step toward prioritizing conservation actions. Our results also suggest that roost specialization may exacerbate population declines due to other risk factors, such as hunting pressure or habitat loss, and thus that management actions to preserve species under risk of extinction should prioritize protection of roosting sites.     

Selectivity in Mammalian Extinction Risk and Threat Types: a New Measure of Phylogenetic Signal Strength in Binary Traits

Abstract: The strength of phylogenetic signal in extinction risk can give insight into the mechanisms behind species’ declines. Nevertheless, no existing measure of phylogenetic pattern in a binary trait, such as extinction‐risk status, measures signal strength in a way that can be compared among data sets. We developed a new measure for phylogenetic signal of binary traits, D, which simulations show gives robust results with data sets of more than 50 species, even when the proportion of threatened species is low. We applied D to the red‐list status of British birds and the world's mammals and found that the threat status for both groups exhibited moderately strong phylogenetic clumping. We also tested the hypothesis that the phylogenetic pattern of species threatened by harvesting will be more strongly clumped than for those species threatened by either habitat loss or invasive species because the life‐history traits mediating the effects of harvesting show strong evolutionary pattern. For mammals, our results supported our hypothesis; there was significant but weaker phylogenetic signal in the risk caused by the other two drivers (habitat loss and invasive species). We conclude that D is likely to be a useful measure of the strength of phylogenetic pattern in many binary traits.     

Regional IUCN Red Listing: the Process as Applied to Birds in the United Kingdom

Abstract:  The World Conservation Union (IUCN) has developed guidelines that enable the assessment of extinction risk at a regional scale. We used these guidelines to assess the extinction risk of birds in the United Kingdom for comparison with an existing assessment of conservation status. Sixty-four species were categorized as regionally threatened, of which 12 were critically endangered. The categorizations of the 223 species assessed agreed broadly with those from the existing U.K. system, which considers more than extinction risk, thus giving a more complete assessment of conservation status. There was, however, a tendency for the IUCN process to give higher risk status to edge-of-range species (some of which are relatively recent colonists considered of comparatively low conservation concern) and low status to those that have declined substantially but remain common (such as many farmland birds, the focus of considerable conservation effort in the United Kingdom). The final red list depended heavily on the subjective decisions made during the assessment process. An alternative interpretation of the guidelines could have resulted in as many as 19 or as few as 6 species being listed as critically endangered. We recommend the revision of the IUCN guidelines to reduce this subjectivity, in particular with respect to the effect of extralimital populations on the likelihood of regional extinction, and hence the potential for variation in the manner of application between regional red-list assessors. Preventing extinction does not have to be the principal driving force behind conservation action at a regional scale if the continuance of a species is safeguarded in other regions.     

Bird Extinctions in Atlantic Forest Fragments of the Viçosa Region, Southeastern Brazil

Abstract:   We studied the conservation status of Atlantic forest birds in 43 forest fragments ranging in size from 1 to 384 ha in the Viçosa region of southeastern Brazil. We compared data from 15 years of field work with historical records from the region, mainly originating from specimens collected by João Moojen during the 1930s. We used published studies associated with museum data and current field work to assess the decline of forest birds during the last 70 years and to relate their disappearance to forest fragmentation and destruction. At least 28 bird species have become locally extinct, 43 are critically endangered, and 25 are vulnerable, representing 60.7% of the original forest bird community known to exist in the region. Vulnerability to fragmentation differed among guilds, forest strata, and endemicity status. Birds that feed on fruit and seeds, and those that feed on insects, were more threatened than omnivores and carnivores. Nectarivorous species were less threatened than other guilds. Moreover, terrestrial and understory birds or birds using only one forest stratum also were more likely to have been threatened. Finally, Atlantic forest endemics were more likely to have become extinct than nonendemic species. In general, sensitivity to environmental disturbance at the local level was similar to the predicted vulnerability to regional disturbance derived from the literature. Our results indicate that a serious decline of Atlantic forest birds is underway and that many other species of birds, not previously recognized as threatened, are of conservation concern .     

The Sensitivity of Population Viability Analysis to Uncertainty about Habitat Requirements: Implications for the Management of the Endangered Southern Brown Bandicoot

Abstract:  Whenever population viability analysis (PVA) models are built to help guide decisions about the management of rare and threatened species, an important component of model building is the specification of a habitat model describing how a species is related to landscape or bioclimatic variables. Model-selection uncertainty may arise because there is often a great deal of ambiguity about which habitat model structure best approximates the true underlying biological processes. The standard approach to incorporate habitat models into PVA is to assume the best habitat model is correct, ignoring habitat-model uncertainty and alternative model structures that may lead to quantitatively different conclusions and management recommendations. Here we provide the first detailed examination of the influence of habitat-model uncertainty on the ranking of management scenarios from a PVA model. We evaluated and ranked 6 management scenarios for the endangered southern brown bandicoot ( Isoodon obesulus ) with PVA models, each derived from plausible competing habitat models developed with logistic regression. The ranking of management scenarios was sensitive to the choice of the habitat model used in PVA predictions. Our results demonstrate the need to incorporate methods into PVA that better account for model uncertainty and highlight the sensitivity of PVA to decisions made during model building. We recommend that researchers search for and consider a range of habitat models when undertaking model-based decision making and suggest that routine sensitivity analyses should be expanded to include an analysis of the impact of habitat-model uncertainty and assumptions.     

Quantitative Determination of Rarity of Freshwater Fishes and Implications for Imperiled‐Species Designations

Abstract: Conserving rare species and protecting biodiversity and ecosystem functioning depends on sound information on the nature of rarity. Rarity is multidimensional and has a variety of definitions, which presents the need for a quantitative classification scheme with which to categorize species as rare or common. We constructed such a classification for North American freshwater fishes to better describe rarity in fishes and provide researchers and managers with a tool to streamline conservation efforts. We used data on range extents, habitat specificities, and local population sizes of North American freshwater fishes and a variety of quantitative methods and statistical decision criteria, including quantile regression and a cost‐function algorithm to determine thresholds for categorizing a species as rare or common. Species fell into eight groups that conform to an established framework for rarity. Fishes listed by the American Fisheries Society (AFS) as endangered, threatened, or vulnerable were most often rare because their local population sizes were low, ranges were small, and they had specific habitat needs, in that order, whereas unlisted species were most often considered common on the basis of these three factors. Species with large ranges generally had few specific habitat needs, whereas those with small ranges tended to have narrow habitat specificities. We identified 30 species not designated as imperiled by AFS that were rare along all dimensions of rarity and may warrant further study or protection, and we found three designated species that were common along all dimensions and may require a review of their imperilment status. Our approach could be applied to other taxa to aid conservation decisions and serve as a useful tool for future revisions of listings of fish species.     

Estimating Extinction Risk with Metapopulation Models of Large‐Scale Fragmentation

Habitat loss is the principal threat to species. How much habitat remains—and how quickly it is shrinking—are implicitly included in the way the International Union for Conservation of Nature determines a species’ risk of extinction. Many endangered species have habitats that are also fragmented to different extents. Thus, ideally, fragmentation should be quantified in a standard way in risk assessments. Although mapping fragmentation from satellite imagery is easy, efficient techniques for relating maps of remaining habitat to extinction risk are few. Purely spatial metrics from landscape ecology are hard to interpret and do not address extinction directly. Spatially explicit metapopulation models link fragmentation to extinction risk, but standard models work only at small scales. Counterintuitively, these models predict that a species in a large, contiguous habitat will fare worse than one in 2 tiny patches. This occurs because although the species in the large, contiguous habitat has a low probability of extinction, recolonization cannot occur if there are no other patches to provide colonists for a rescue effect. For 4 ecologically comparable bird species of the North Central American highland forests, we devised metapopulation models with area‐weighted self‐colonization terms; this reflected repopulation of a patch from a remnant of individuals that survived an adverse event. Use of this term gives extra weight to a patch in its own rescue effect. Species assigned least risk status were comparable in long‐term extinction risk with those ranked as threatened. This finding suggests that fragmentation has had a substantial negative effect on them that is not accounted for in their Red List category. Estimación del Riesgo de Extinción Mediante Modelos Metapoblacionales de Fragmentación a Gran Escala     

Global Warming, Human Population Pressure, and Viability of the World's Smallest Butterfly

Abstract:  The effects of climate change and habitat destruction and their interaction are likely to be the greatest challenge to animal and plant conservation in the twenty-first century. We used the world's smallest butterfly, the Sinai baton blue ( Pseudophilotes sinaicus ), as an exemplar of how global warming and human population pressures may act together to cause species extinctions. We mapped the entire global range of this butterfly and obtained extensive data on the intensity of livestock grazing. As with an increasing number of species, it is confined to a network of small habitat patches and is threatened both by indirect human-induced factors (global warming) and by the direct activities of humans (in this case, livestock grazing and collection of medicinal plants). In the absence of global warming, grazing, and plant collection, our model suggested that the butterfly will persist for at least 200 years. Above a threshold intensity of global warming, the chance of extinction accelerated rapidly, implying that there may be an annual average temperature, specific to each endangered species, above which extinction becomes very much more likely. By contrast, there was no such threshold of grazing pressure—the chance of extinction increased steadily with increasing grazing. The impact of grazing, however, decreased with higher levels of year-to-year variation in habitat quality. The effect of global warming did not depend on the future level of grazing, suggesting that the impacts of global warming and grazing are additive. If the areas of habitat patches individually fall below certain prescribed levels, the butterfly is likely to go extinct. Two patches were very important for persistence: if either were lost the species would probably go extinct. Our results have implications for the conservation management of all species whose habitats are at risk because of the direct activities of humans and in the longer term because of climate change.     

Effects of Human Activity on Global Extinction Risk

Both natural and anthropogenic factors are important in determining a species' risk of extinction. Little work has been done, however, to quantify the magnitude of current anthropogenic influences on the extinction process. The purpose of this study is to determine the extent to which measures of the intensity of human activity are related to the global variability of two measures of species' susceptibility to extinction. We observed six indices of human activities in 90 countries, and we tested their relationships to the proportion of threatened bird and mammal species in each country, as well as to mammalian population density. After correcting for area effects, latitudinal diversity gradients, and body size (for population density), 28 to 50% of the remaining variation was statistically attributable to anthropogenic variables. Different measures of anthropogenic influence were most closely related to extinction risk in birds and mammals. Human population density was the variable most closely related to the proportion of threatened bird species per country, whereas per capita GNP was more important for mammal species. Mammalian population density strongly correlates with the extent of protected area per country. Contrary to suggestions in earlier literature, our work does not support the hypothesis that habitat loss is a prime contributor to species loss because frequencies of threatened birds and mammals are not closely related to patterns of land use.     

The importance of incorporating functional habitats into conservation planning for highly mobile species in dynamic systems

The distribution of mobile species in dynamic systems can vary greatly over time and space. Estimating their population size and geographic range can be problematic and affect the accuracy of conservation assessments. Scarce data on mobile species and the resources they need can also limit the type of analytical approaches available to derive such estimates. We quantified change in availability and use of key ecological resources required for breeding for a critically endangered nomadic habitat specialist, the Swift Parrot (Lathamus discolor). We compared estimates of occupied habitat derived from dynamic presence‐background (i.e., presence‐only data) climatic models with estimates derived from dynamic occupancy models that included a direct measure of food availability. We then compared estimates that incorporate fine‐resolution spatial data on the availability of key ecological resources (i.e., functional habitats) with more common approaches that focus on broader climatic suitability or vegetation cover (due to the absence of fine‐resolution data). The occupancy models produced significantly (P < 0.001) smaller (up to an order of magnitude) and more spatially discrete estimates of the total occupied area than climate‐based models. The spatial location and extent of the total area occupied with the occupancy models was highly variable between years (131 and 1498 km²). Estimates accounting for the area of functional habitats were significantly smaller (2–58% [SD 16]) than estimates based only on the total area occupied. An increase or decrease in the area of one functional habitat (foraging or nesting) did not necessarily correspond to an increase or decrease in the other. Thus, an increase in the extent of occupied area may not equate to improved habitat quality or function. We argue these patterns are typical for mobile resource specialists but often go unnoticed because of limited data over relevant spatial and temporal scales and lack of spatial data on the availability of key resources. Understanding changes in the relative availability of functional habitats is crucial to informing conservation planning and accurately assessing extinction risk for mobile resource specialists.     

Bird Conservation in Brazil

Abstract:  Brazil has one of the richest avifaunas in the world, with recent estimates varying from 1696 to 1731 species. About 10% (193 taxa) of these are threatened. The Amazon has the highest number of species, followed by the Atlantic Forest and the Cerrado; most of Brazil's endemic birds, however, are in the Atlantic Forest. Brazil's threatened species occur mostly in the Atlantic Forest, especially in the southeast lowlands and the northeast. The Cerrado has the second highest number of threatened species. The two major threats to Brazilian birds are habitat loss, degradation, and fragmentation and hunting, most especially for illegal commerce. A number of conservation and research initiatives over the last 20 years have significantly improved our capacity to address and resolve major issues for bird conservation. Brazil requires a National Bird Conservation Plan to draw up priorities for research and conservation over the next decade.     

Landscape Characteristics of Fragmented Shrubsteppe Habitats and Breeding Passerine Birds

We examined the influence of local and landscape-level attributes of fragmented habitats in shrubsteppe habitats on the breeding distributions of Sage ( Amphispiza belli ) and Brewer's ( Spizella breweri ) Sparrows, Sage Thrashers ( Oreoscoptes montanus ) Horned Larks ( Eremophila alpestris ), and Western Meadowlarks ( Sturnella neglecta ) in the Snake River Plains of southwestern Idaho. We developed habitat (resource) selection models for each species by combining bird counts conducted from 1991 through 1933 with local vegetation characteristics and landscape attributes derived from satellite imagery. Site selection by shrubsteppe species (Sage and Brewer's Sparrows, and Sage Thrashers) depended on local vegetation cover and landscape features, such as the patch size of shrub habitats or the spatial similarity of sites. Marginal sites for these species (with species present in one of three years) were intermediate between unoccupied (never present) and occupied sites along environmental gradients characterized by increasing size of shrub habitat patches and total shrub cover and by decreasing disturbance. Horned Larks and Western Meadowlarks, typical grassland species, were not sensitive to landscape features, and their occupancy depended on the amount of grassland or shrub cover. In contrast to shrubsteppe species, sites that varied by occupancy rates of Western Meadowlarks did not significantly differ in vegetation or landscape components. Our results demonstrate that fragmentation of shrubsteppe significantly influenced the presence of shrub-obligate species. Because of restoration difficulties, the disturbance of semiarid shrubsteppe may cause irreversible loss of habitat and significant long-term consequences for the conservation of shrub-obligate birds.     

Bird Species' Tolerance of Secondary Forest Habitats and Its Effects on Extinction

Abstract:  Intense deforestation causes massive species losses. These losses occur because the habitats supplanting primary forest are inadequate to sustain viable populations of forest-dependent species. Despite this, certain species do seem to persist within the secondary habitats that replace original forest. This implies that there is a special class of species that might survive the loss of primary forest. Such a result would significantly influence conservation plans and extinction predictions. We tested whether species that tolerate secondary habitats survive extensive habitat loss and whether the same degree of loss threatens species that are forest obligates. To identify purported "survivors," we compared the remaining range sizes of endemic birds, their abundances, and their degree of extinction threat. We did this within the remaining Atlantic Forest of Brazil, a region extremely rich in endemics but with only approximately 10% of its forest remaining. We found no survivors. Habitat loss threatens forest-obligate birds and those using secondary habitats equally.     

Effects of anthropogenic landscapes on population maintenance of waterbirds

Anthropogenic impacts have reduced natural areas but increased the area of anthropogenic landscapes. There is debate about whether anthropogenic landscapes (e.g., farmlands, orchards, and fish ponds) provide alternatives to natural habitat and under what circumstances. We considered whether anthropogenic landscapes can mitigate population declines for waterbirds. We collected data on population trends and biological traits of 1203 populations of 579 species across the planet. Using Bayesian generalized linear mixed models, we tested whether the ability of a species to use an anthropogenic landscape can predict population trends of waterbird globally and of species of conservation concern. Anthropogenic landscapes benefited population maintenance of common but not less-common species. Conversely, the use of anthropogenic landscapes was associated with population declines for threatened species. Our findings delineate some limitations to the ability of anthropogenic landscapes to mitigate population declines, suggesting that the maintenance of global waterbird populations depends on protecting remaining natural areas and improving the habitat quality in anthropogenic landscapes. Article impact statement: Protecting natural areas and improving the quality of anthropogenic landscapes as habitat are both needed to achieve effective conservation.     

Environmental Synergisms and Extinctions of Tropical Species

Abstract: Environmental synergisms may pose the greatest threat to tropical biodiversity. Using recently updated data sets from the International Union for Conservation of Nature (IUCN) Red List, we evaluated the incidence of perceived threats to all known mammal, bird, and amphibian species in tropical forests. Vulnerable, endangered, and extinct species were collectively far more likely to be imperiled by combinations of threats than expected by chance. Among 45 possible pairwise combinations of 10 different threats, 69%, 93%, and 71% were significantly more frequent than expected for threatened mammals, birds, and amphibians, respectively, even with a stringent Bonferroni‐corrected probability value (p= 0.003). Based on this analysis, we identified five key environmental synergisms in the tropics and speculate on the existence of others. The most important involve interactions between habitat loss or alteration (from agriculture, urban sprawl, infrastructure, or logging) and other anthropogenic disturbances such as hunting, fire, exotic‐species invasions, or pollution. Climatic change and emerging pathogens also can interact with other threats. We assert that environmental synergisms are more likely the norm than the exception for threatened species and ecosystems, can vary markedly in nature among geographic regions and taxa, and may be exceedingly difficult to predict in terms of their ultimate impacts. The perils posed by environmental synergisms highlight the need for a precautionary approach to tropical biodiversity conservation.