From Red Lists to Species of Conservation Concern

Abstract:  National red lists of threatened animal and plant species prepared according to the criteria of the World Conservation Union (IUCN) adequately reflect the extinction risk of species within a country but cannot be used directly to set conservation priorities. In particular, the significance of national populations for the conservation of the species as a whole is not taken into account. We present a procedure that can be used to assess national responsibility based on the national red-list status of a species, the international importance of the national population, and the species' "historical rarity" status. We distinguished five responsibility classes for breeding birds: B1, threatened species with internationally important populations in Switzerland; B2, threatened species with internationally less important populations; B3, nonthreatened species with internationally important populations; B4, nonthreatened species with internationally less important populations; and B5, species that have never been common in Switzerland. Two responsibility classes were distinguished for birds occurring in Switzerland as visitors: G1, species with large concentrations in Switzerland and an unfavorable conservation status in Europe, and G2, species with large concentrations in Switzerland and a favorable conservation status in Europe. Two additional classes (G3 and G4) for visiting species occurring in internationally less important numbers are possible but were not analyzed in detail. Responsibility classes B1, B2, B3, G1, and G2 were defined as species of national conservation concern. We developed the method for birds in Switzerland, but it can be used in other countries and for other taxonomic groups as well. It is particularly suitable where national red lists are established according to IUCN guidelines.     

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.     

Conserving the abundance of nonthreatened species

Human modification of the environment is driving declines in population size and distributional extent of much of the world's biota. These declines extend to many of the most abundant and widespread species, for which proportionally small declines can result in the loss of vast numbers of individuals, biomass, and interactions. These losses could have major localized effects on ecological and cultural processes and services without elevating a species’ global extinction risk. Although most conservation effort is directed at species threatened with extinction in the very near term, the value of retaining abundance regardless of global extinction risk is justifiable based on many biodiversity or ecosystem service metrics, including cultural services, at scales from local to global. The challenges of identifying conservation priorities for widespread and abundant species include quantifying the effects of species’ abundance on services and understanding how these effects are realized as populations decline. Negative effects of population declines may be disconnected from the threat processes driving declines because of species movements and environment flows (e.g., hydrology). Conservation prioritization for these species shares greater similarity with invasive species risk assessments than extinction risk assessments because of the importance of local context and per capita effects of abundance on other species. Because conservation priorities usually focus on preventing the extinction of threatened species, the rationale and objectives for incorporating declines of nonthreatened species must be clearly articulated, going beyond extinction risk to encompass the range of likely harmful effects (e.g., secondary extinctions, loss of ecosystem services) if declines persist or are not reversed. Research should focus on characterizing the effects of local declines in species that are not threatened globally across a range of ecosystem services and quantifying the spatial distribution of these effects through the distribution of abundance. The case for conserving abundance in nonthreatened species can be made most powerfully when the costs of losing this abundance are better understood.     

China's endemic vertebrates sheltering under the protective umbrella of the giant panda

The giant panda attracts disproportionate conservation resources. How well does this emphasis protect other endemic species? Detailed data on geographical ranges are not available for plants or invertebrates, so we restrict our analyses to 3 vertebrate taxa: birds, mammals, and amphibians. There are gaps in their protection, and we recommend practical actions to fill them. We identified patterns of species richness, then identified which species are endemic to China, and then which, like the panda, live in forests. After refining each species' range by its known elevational range and remaining forest habitats as determined from remote sensing, we identified the top 5% richest areas as the centers of endemism. Southern mountains, especially the eastern Hengduan Mountains, were centers for all 3 taxa. Over 96% of the panda habitat overlapped the endemic centers. Thus, investing in almost any panda habitat will benefit many other endemics. Existing panda national nature reserves cover all but one of the endemic species that overlap with the panda's distribution. Of particular interest are 14 mammal, 20 bird, and 82 amphibian species that are inadequately protected. Most of these species the International Union for Conservation of Nature currently deems threatened. But 7 mammal, 3 bird, and 20 amphibian species are currently nonthreatened, yet their geographical ranges are <20,000 km² after accounting for elevational restriction and remaining habitats. These species concentrate mainly in Sichuan, Yunnan, Nan Mountains, and Hainan. There is a high concentration in the east Daxiang and Xiaoxiang Mountains of Sichuan, where pandas are absent and where there are no national nature reserves. The others concentrate in Yunnan, Nan Mountains, and Hainan. Here, 10 prefectures might establish new protected areas or upgrade local nature reserves to national status.     

Incorporating Evolutionary Measures into Conservation Prioritization

Abstract:  Conservation prioritization is dominated by the threat status of candidate species. However, species differ markedly in the shared genetic information they embody, and this information is not taken into account if species are prioritized by threat status alone. We developed a system of prioritization that incorporates both threat status and genetic information and applied it to 9546 species of birds worldwide. We devised a simple measure of a species' genetic value that takes into account the shape of the entire taxonomic tree of birds. This measure approximates the evolutionary history that each species embodies and sums to the phylogenetic diversity of the entire taxonomic tree. We then combined this genetic value with each species' probability of extinction to create a species-specific measure of expected loss of genetic information. The application of our methods to the world's avifauna showed that ranking species by expected loss of genetic information may help preserve bird evolutionary history by upgrading those threatened species with fewer close relatives. We recommend developing a mechanism to incorporate a species' genetic value into the prioritization framework.     

Climate Change, Elevational Range Shifts, and Bird Extinctions

Abstract:  Limitations imposed on species ranges by the climatic, ecological, and physiological effects of elevation are important determinants of extinction risk. We modeled the effects of elevational limits on the extinction risk of landbirds, 87% of all bird species. Elevational limitation of range size explained 97% of the variation in the probability of being in a World Conservation Union category of extinction risk. Our model that combined elevational ranges, four Millennium Assessment habitat-loss scenarios, and an intermediate estimate of surface warming of 2.8° C, projected a best guess of 400–550 landbird extinctions, and that approximately 2150 additional species would be at risk of extinction by 2100. For Western Hemisphere landbirds, intermediate extinction estimates based on climate-induced changes in actual distributions ranged from 1.3% (1.1° C warming) to 30.0% (6.4° C warming) of these species. Worldwide, every degree of warming projected a nonlinear increase in bird extinctions of about 100–500 species. Only 21% of the species predicted to become extinct in our scenarios are currently considered threatened with extinction. Different habitat-loss and surface-warming scenarios predicted substantially different futures for landbird species. To improve the precision of climate-induced extinction estimates, there is an urgent need for high-resolution measurements of shifts in the elevational ranges of species. Given the accelerating influence of climate change on species distributions and conservation, using elevational limits in a tested, standardized, and robust manner can improve conservation assessments of terrestrial species and will help identify species that are most vulnerable to global climate change. Our climate-induced extinction estimates are broadly similar to those of bird species at risk from other factors, but these estimates largely involve different sets of species.     

Species Inequality in Scientific Study

Abstract: Some conservationists argue for a focused effort to protect the most critically endangered species, and others suggest a large‐scale endeavor to safeguard common species across large areas. Similar arguments are applicable to the distribution of scientific effort among species. Should conservation scientists focus research efforts on threatened species, common species, or do all species deserve equal attention? We assessed the scientific equity among 1909 mammals, birds, reptiles, and amphibians of southern Africa by relating the number of papers written about each species to their status on the International Union for Conservation of Nature Red List. Threatened large mammals and reptiles had more papers written about them than their nonthreatened counterparts, whereas threatened small mammals and amphibians received less attention than nonthreatened species. Threatened birds received an intermediate amount of attention in the scientific literature. Thus, threat status appears to drive scientific effort among some animal groups, whereas other factors (e.g., pest management and commercial interest) appear to dictate scientific investment in particular species of other groups. Furthermore, the scientific investment per species differed greatly between groups—the mean number of papers per threatened large mammal eclipsed that of threatened reptiles, birds, small mammals, and amphibians by 2.6‐, 15‐, 216‐, and more than 500‐fold, respectively. Thus, in the eyes of science, all species are not created equal. A few species commanded a great proportion of scientific attention, whereas for many species information that might inform conservation is virtually nonexistent.     

Impact of alternative metrics on estimates of extent of occurrence for extinction risk assessment

In International Union for Conservation of Nature (IUCN) Red List assessments, extent of occurrence (EOO) is a key measure of extinction risk. However, the way assessors estimate EOO from maps of species’ distributions is inconsistent among assessments of different species and among major taxonomic groups. Assessors often estimate EOO from the area of mapped distribution, but these maps often exclude areas that are not habitat in idiosyncratic ways and are not created at the same spatial resolutions. We assessed the impact on extinction risk categories of applying different methods (minimum convex polygon, alpha hull) for estimating EOO for 21,763 species of mammals, birds, and amphibians. Overall, the percentage of threatened species requiring down listing to a lower category of threat (taking into account other Red List criteria under which they qualified) spanned 11–13% for all species combined (14–15% for mammals, 7–8% for birds, and 12–15% for amphibians). These down listings resulted from larger estimates of EOO and depended on the EOO calculation method. Using birds as an example, we found that 14% of threatened and near threatened species could require down listing based on the minimum convex polygon (MCP) approach, an approach that is now recommended by IUCN. Other metrics (such as alpha hull) had marginally smaller impacts. Our results suggest that uniformly applying the MCP approach may lead to a one‐time down listing of hundreds of species but ultimately ensure consistency across assessments and realign the calculation of EOO with the theoretical basis on which the metric was founded.     

Identifying and Managing Threatened Invertebrates through Assessment of Coextinction Risk

Abstract: Invertebrates with specific host species may have a high probability of extinction when their hosts have a high probability of extinction. Some of these invertebrates are more likely to go extinct than their hosts, and under some circumstances, specific actions to conserve the host may be detrimental to the invertebrate. A critical constraint to identifying such invertebrates is uncertainty about their level of host specificity. We used two host‐breadth models that explicitly incorporated uncertainty in the host specificity of an invertebrate species. We devised a decision protocol to identify actions that may increase the probability of persistence of a given dependent species. The protocol included estimates from the host‐breadth models and decision nodes to identify cothreatened species. We applied the models and protocol to data on 1055 insects (186 species) associated with 2 threatened (as designated by the Australian Government) plant species and 19 plant species that are not threatened to determine whether any insect herbivores have the potential to become extinct if the plant becomes extinct. According to the host‐breadth models, 18 species of insect had high host specificity to the threatened plant species. From these 18 insects, the decision protocol highlighted 6 species that had a high probability of extinction if their hosts were to become extinct (3% of all insects examined). The models and decision protocol have added objectivity and rigor to the process of deciding which dependent invertebrates require conservation action, particularly when dealing with largely unknown and speciose faunas.     

Geographic Range Fragmentation and Abundance in Neotropical Migratory Birds

Populations of neotropical migrant landbirds have experienced significant declines in recent years. We investigated potential consequences of these declines by examining the relationship between abundance and fragmentation of geographic ranges of species on the North American breeding grounds. We estimated areograpbic fragmentation using the box dimension of a species' geographic range and demographic fragmentation using the fractal dimension of the semivariance function calculated from samples of population abundance across species' geographic ranges. We found a negative relationship between average abundance and demographic fragmentation for neotropical migrants, but not for residents. We also showed that demographic fragmentation and areographic fragmentation are inversely related for residents, but not for neotropical migrants. These results imply that neotropical migrants may be more sensitive to extinction than are residents.     

A Mechanistic Approach to Evaluation of Umbrella Species as Conservation Surrogates

Abstract:  Although species with large area requirements are sometimes used as umbrella species, their general utility as conservation tools is uncertain. We surveyed the species diversity of birds, butterflies, carabids, and forest-floor plants in forest sites across an area (1600 km²) in which we delineated large breeding home ranges of Northern Goshawk ( Accipiter gentilis ). We tested whether protection of the home ranges could serve as an effective umbrella to protect sympatric species of the four taxa. We also used an empirical habitat model of occupancy of home range to examine mechanisms by which the Northern Goshawk acts as an umbrella species. Among species richness, abundance, and species composition of the four taxa, only abundance and species composition of birds differed between sites located inside and outside home ranges, which was due to greater abundance of bird species that were prey of Northern Goshawks inside the home ranges. Thus, although home range indicated areas with high abundance of certain bird prey species, it was not effective as an indicator of the species diversity of all four taxa. We also did not find any difference in species richness, abundance, and species composition between sites predicted as occupied and unoccupied using the habitat model. In contrast, when we selected sites on the basis of each habitat variable in the model, habitat variables that selected sites either in agricultural or forested landscapes encompassed sites with high species richness or particular species composition. This result suggests that the low performance of the Northern Goshawk as an umbrella species is due to this species' preference for habitat in both agricultural and forested landscapes. Species that can adjust to changes in habitat conditions may not act as effective umbrella species despite having large home ranges.     

The Bias of Complementarity Hotspots toward Marginal Populations

Abstract: It has been suggested that using complementarity to identify networks of important areas for conserving biodiversity may preferentially select areas within the margins of species ranges. We tested this idea by examining the location of complementarity hotspots in relation to two measures of range core-periphery. The first measures patterns of aggregation among records within each species' range to identify areas within the core (i.e., areas with aggregated distributions) and periphery (i.e., areas with scattered distributions) of the range. The second measures spatial turnover among species to identify areas with a high density of range edges. For three selected groups of terrestrial vertebrates in Europe—mammals, birds, and herptiles—areas chosen based on complementarity were located within the margins of species' ranges more often than expected by chance. This pattern was consistent for the two measures of core-periphery we used. The bias of complementarity hotspots toward marginal populations is especially important for species with restricted range sizes. If extinctions are determined mainly by demographic factors, then selecting areas at the peripheries of species' ranges might be a poor option. But if extinctions are determined mainly by extrinsic factors, then peripheral populations might be important to ensure the long-term persistence of species.     

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.     

The Normative Dimension and Legal Meaning of Endangered and Recovery in the U.S. Endangered Species Act

Abstract:  The ethical, legal, and social significance of the U.S. Endangered Species Act of 1973 (ESA) is widely appreciated. Much of the significance of the act arises from the legal definitions that the act provides for the terms threatened species and endangered species. The meanings of these terms are important because they give legal meaning to the concept of a recovered species. Unfortunately, the meanings of these terms are often misapprehended and rarely subjected to formal analysis. We analyzed the legal meaning of recovered species and illustrate key points with details from "recovery" efforts for the gray wolf ( Canis lupus ). We focused on interpreting the phrase "significant portion of its range," which is part of the legal definition of endangered species. We argue that recovery and endangerment entail a fundamentally normative dimension (i.e., specifying conditions of endangerment) and a fundamentally scientific dimension (i.e., determining whether a species meets the conditions of endangerment). Specifying conditions for endangerment is largely normative because it judges risks of extinction to be either acceptable or unacceptable. Like many other laws that specify what is unacceptable, the ESA largely specifies the conditions that constitute unacceptable extinction risk. The ESA specifies unacceptable risks of extinction by defining endangered species in terms of the portion of a species' range over which a species is "in danger of extinction." Our analysis indicated that (1) legal recovery entails much more than the scientific notion of population viability, (2) most efforts to recover endangered species are grossly inadequate, and (3) many unlisted species meet the legal definition of an endangered or threatened species.     

Identifying conservation priorities for threatened Eastern Himalayan mammals

To augment mammal conservation in the Eastern Himalayan region, we assessed the resident 255 terrestrial mammal species and identified the 50 most threatened species based on conservation status, endemism, range size, and evolutionary distinctiveness. By using the spatial analysis package letsR and the complementarity core‐area method in the conservation planning software Zonation, we assessed the current efficacy of their protection and identified priority conservation areas by comparing protected areas (PAs), land cover, and global ecoregion 2017 maps at a 100 × 100 m spatial scale. The 50 species that were most threatened, geographically restricted, and evolutionarily distinct faced a greater extinction risk than globally nonthreatened and wide‐ranging species and species with several close relatives. Small, medium‐sized, and data‐deficient species faced extinction from inadequate protection in PAs relative to wide‐ranging charismatic species. There was a mismatch between current PA distribution and priority areas for conservation of the 50 most endangered species. To protect these species, the skewed regional PA distribution would require expansion. Where possible, new PAs and transboundary reserves in the 35 priority areas we identified should be established. There are adequate remaining natural areas in which to expand current Eastern Himalayan PAs. Consolidation and expansion of PAs in the EH requires strengthening national and regional transboundary collaboration, formulating comprehensive regional land‐use plans, diversifying conservation funding, and enhancing information sharing through a consolidated regional database.     

Phylogenetic Comparative Methods Strengthen Evidence for Reduced Genetic Diversity among Endangered Tetrapods

Abstract: The fitness of species with little genetic diversity is expected to be affected by inbreeding and an inability to respond to environmental change. Conservation theory suggests that endangered species will generally demonstrate lower genetic diversity than taxa that are not threatened. This hypothesis has been challenged because the time frame of anthropogenic extinction may be too fast to expect genetic factors to significantly contribute. I conducted a meta‐analysis to examine how genetic diversity in 894 tetrapods correlates with extinction threat level. Because species are not evolutionarily independent, I used a phylogenetic regression framework to address this issue. Mean genetic diversity of tetrapods, as assessed by protein heterozygosity, was 29.7–31.5% lower on average in threatened species than in their nonthreatened relatives, a highly significant reduction. Within amphibians as diversity decreased extinction risk increased in phylogenetic models, but not in nonphylogenetic regressions. The effects of threatened status on diversity also remained significant after accounting for body size in mammals. These results support the hypothesis that genetic effects on population fitness are important in the extinction process.     

Effects of Economic Prosperity on Numbers of Threatened Species

Abstract: We used data from over 100 countries to investigate the link between numbers of threatened species and per-capita gross national product. We corrected for factors that might otherwise confound such a relationship. Our study was motivated by the continuing debate over the relationship between environmental degradation and per-capita income. Proponents of the environmental Kuznets-curve hypothesis argue that although environmental degradation may increase initially, increases in per-capita income will eventually result in greater environmental quality. Theoretical objections and the lack of widespread empirical evidence recently have thrown doubt on the existence of such a pattern. Treating threat to biodiversity as one potential indicator of environmental degradation, we divided threatened species into seven taxonomic groups ( plants, mammals, birds, amphibians, reptiles, fishes, and invertebrates) and analyzed each group separately. Count-data regression analysis indicated that the number of threatened species was related to per-capita gross national product in five of seven taxonomic groups. Birds were the only taxonomic group in which numbers of threatened species decreased throughout the range of developed countries' per-capita gross national product. Plants, amphibians, reptiles, and invertebrates showed increasing numbers of threatened species throughout this same range. If these relationships hold, increasing numbers of species from several taxonomic groups are likely to be threatened with extinction as countries increase in prosperity. A key challenge is to understand the interactions among consumer preferences, biology, and institutions that lead to the relationship observed for birds and to see whether this knowledge can be applied to conservation of other taxa.     

A composite measure of habitat loss for entire assemblages of species

Habitat destruction is among the greatest threats facing biodiversity, and it affects common and threatened species alike. However, metrics for communicating its impacts typically overlook the nonthreatened component of assemblages. This risks the loss of habitat going unreported for species that comprise the majority of assemblages. We adapted a widely used measure for summarizing researcher output (the h index) to provide a metric that describes natural habitat loss for entire assemblages, inclusive of threatened and nonthreatened species. For each of 447 Australian native terrestrial bird species, we combined information on their association with broad vegetation groups with distributional range maps to identify the difference between the estimated pre-European and current extents of potential habitat, defined as vegetation groups most closely associated with each species. From this, we calculated the loss index (LI), which revealed that 30% of native birds have each lost at least 30% of their potential natural habitat (LI = 30). At the subcontinental scale, LIs ranged from 15 in arid Australia to 61 in the highly transformed southeastern part of the country. Different subcomponents of the assemblage had different LI values. For example, Australia's parrots (n = 52 species) had an LI of 38, whereas raptors (n = 32 species) had an LI of 25. The LI is simple to calculate and can be determined using readily available spatial information on species distributions, native vegetation associations, and human impacts on natural land cover. This metric, including the curves used to deduce it, could complement other biodiversity indices if it is used for regional and global biodiversity assessments that compare the status of natural habitat extent for assemblages within and among nations, monitor changes through time, and forecast future changes to guide strategic land-use planning. The LI is an intuitive tool that can be used to summarize and communicate how human actions affect whole assemblages, not just threatened species.     

Vulnerability and Resilience of Tropical Forest Species to Land‐Use Change

Abstract: We provide a cross‐taxon and historical analysis of what makes tropical forest species vulnerable to extinction. Several traits have been important for species survival in the recent and distant geological past, including seed dormancy and vegetative growth in plants, small body size in mammals, and vagility in insects. For major past catastrophes, such as the five mass extinction events, large range size and vagility or dispersal were key to species survival. Traits that make some species more vulnerable to extinction are consistent across time scales. Terrestrial organisms, particularly animals, are more extinction prone than marine organisms. Plants that persist through dramatic changes often reproduce vegetatively and possess mechanisms of die back. Synergistic interactions between current anthropogenic threats, such as logging, fire, hunting, pests and diseases, and climate change are frequent. Rising temperatures threaten all organisms, perhaps particularly tropical organisms adapted to small temperature ranges and isolated by distance from suitable future climates. Mutualist species and trophic specialists may also be more threatened because of such range‐shift gaps. Phylogenetically specialized groups may be collectively more prone to extinction than generalists. Characterization of tropical forest species’ vulnerability to anthropogenic change is constrained by complex interactions among threats and by both taxonomic and ecological impediments, including gross undersampling of biotas and poor understanding of the spatial patterns of taxa at all scales.     

Region-wide retreats from lower elevations of range-restricted birds across the Northern Andes

Local studies show upslope shifts in the distribution of tropical birds in response to warming temperatures. Unanswered is whether these upward shifts occur regionally across many species. We considered a nearly 2000-km length of the Northern Andes, where deforestation, temperature, and extreme weather events have increased during the past decades. Range-restricted bird species are particularly vulnerable to such events and occur in exceptionally high numbers in this region. Using abundant crowd-sourced data from the Cornell Lab of Ornithology database, eBird, and the Global Biodiversity Information Facility, we documented distributions of nearly 200 such species. We examined whether species shifted their elevational ranges over time by comparing observed versus expected occurrences below a low elevational threshold and above a high elevational threshold for 2 periods: before and after 2005. We predicted fewer observations at lower elevations (those below the threshold) and more at upper elevations (those above the threshold) after 2005. We also tested for deforestation effects at lower elevations within each species’ distribution ranges. We compared relative forest loss with the differences between observed and expected occurrences across the elevational range. Species’ retreats from lower elevations were ubiquitous and involved a 23–40% decline in prevalence at the lowest elevations. Increases at higher elevations were not consistent. The retreats occurred across a broad spectrum of species, from predominantly lowland to predominantly highland. Because deforestation showed no relationship with species retreats, we contend that a warming climate is the most parsimonious explanation for such shifts.