Interspecific Patterns of Genetic Diversity in Birds: Correlations with Extinction Risk

Abstract:  Birds are frequently used as indicators of ecosystem health and are the most comprehensively studied class in the animal kingdom. Nevertheless, a comprehensive, interspecific assessment of the correlates of avian genetic diversity is lacking, even though indices of genetic diversity are of considerable interest in the conservation of threatened species. We used published data on variation at microsatellite loci from 194 bird species to examine correlates of diversity, particularly with respect to conservation status and population size. We found a significant decline in mean heterozygosity with increasing extinction risk, and showed, by excluding species whose heterozygosity values were calculated with heterospecific primers, that this relationship was not dependent on ascertainment bias. Results of subsequent regression analyses suggested that smaller population sizes of threatened species were largely responsible for this relationship. Thus, bird species at risk of extinction are relatively depauperate in terms of neutral genetic diversity, which is expected to make population recovery more difficult if it reflects adaptive genetic variation. Conservation policy will need to minimize further loss of diversity if the chances of saving threatened species are to be maximized.     

A Reassessment of Homozygosity and the Case for Inbreeding Depression in the Cheetah, Acinonyx jubatus: Implications for Conservation

Preservation of genetic diversity within declining populations of endangered species is a major concern in the discipline of conservation biology. The endangered cheetah, Acinonyx jubatus , exhibits relatively little genetic variability (polymorphism = 0.02–0.04, heterozygosity = 0.0004–0.014). Since the discovery of the cheetah's relative homozygosity, this species has been frequently cited as an example of one whose survival may be compromised by the loss of genetic diversity. The cheetah's genetic uniformity is generally believed to be the result of an historical population bottle-neck followed by a high level of inbreeding. Evidence offered in support of this hypothesis includes the cheetah's present low level of genetic variability and symptoms of inbreeding depression in captive populations. Using available data on fluctuating asymmetry and genetic variation in other carnivores, I question the assumption that the present level of genetic diversity in the cheetah is indicative of a loss of former variability. Carnivores exhibit significantly lower levels of genetic variation than other mammals, and several carnivores for which data are available exhibit lower levels of heterozygosity and polymorphism than the cheetah does. Measures of fluctuating asymmetry do not support the hypothesis that the cheetah is suffering an increased level of bomozygosity due to genetic stress. Many of the phenotypic effects attributed to inbreeding depression, such as infertility, reduced litter sizes, and increased susceptibility to disease, are limited to captive individuals and may be explained as physiological or behavioral artifacts of captivity. In sum, the genetic constitution of the cheetah does not appear to compromise the survival of the species. Conservation efforts may be more effectively aimed at a real, immediate threat to the cheetah's future: the loss of its natural habitat.     

Genetic structure,lack of sex-biased dispersal and behavioral flexibility in the pair-living fat-tailed dwarf lemur, <Emphasis Type="Italic">Cheirogaleus medius</Emphasis>

Mating system and dispersal patterns influence the spatio-genetic structure within and between populations. Among mammals, monogamy is rare, and its socio-genetic consequences have not been studied in detail before. The goal of our study was to investigate population history, demographic structure, and dispersal patterns in a population of pair-living fat-tailed dwarf lemurs, Cheirogaleus medius, a small, nocturnal primate from western Madagascar, and to infer their underlying behavioral mechanisms. Tissue samples for DNA extraction were obtained from a total of 140 individuals that were captured in two subpopulations about 3 km apart. Analyses of mtDNA variability at the population level revealed very low levels of genetic variability combined with high haplotype diversity, which is indicative of a recent population bottleneck. We found no evidence for spatial clustering of same-sexed individuals with identical haplotypes within each of two subpopulations but significant clustering between them. Thus, a high level of local subpopulation differentiation was observed (F _ST = 0.230). The sexes showed equal variances in the number of individuals representing each haplotype, as well as equal levels of aggregation of identical haplotypes. Hence, both sexes disperse from their natal area, one pattern expected in a pair-living mammal. There is a possibility of behavioral and social flexibility in this species, however, because we documented pronounced differences in density and sex ratio between the two subpopulations, suggesting that single study sites or populations may not be representative of a given local population or even species.     

Genetic Bottlenecks Driven by Population Disconnection

Abstract: Connectivity among populations plays a crucial role in maintaining genetic variation at a local scale, especially in small populations affected strongly by genetic drift. The negative consequences of population disconnection on allelic richness and gene diversity (heterozygosity) are well recognized and empirically established. It is not well recognized, however, that a sudden drop in local effective population size induced by such disconnection produces a temporary disequilibrium in allelic frequency distributions that is akin to the genetic signature of a demographic bottleneck. To document this effect, we used individual‐based simulations and empirical data on allelic richness and gene diversity in six pairs of isolated versus well‐connected (core) populations of European tree frogs. In our simulations, population disconnection depressed allelic richness more than heterozygosity and thus resulted in a temporary excess in gene diversity relative to mutation drift equilibrium (i.e., signature of a genetic bottleneck). We observed a similar excess in gene diversity in isolated populations of tree frogs. Our results show that population disconnection can create a genetic bottleneck in the absence of demographic collapse.     

Translocations and the Preservation of Allelic Diversity

Translocation is a tool commonly used for the conservation of threatened and endangered fish species. Despite extensive use, the biological implications of translocation remain poorly understood. Of particular interest is the effect of translocation on genetic variability. Maintenance of genetic variability in these "refuge" populations is assumed to be important for both short- and long-term success. We examined allozyme variability at 16 loci for western mosquitofish ( Gambusia affinis ) populations with known histories of introduction. Refuge populations had significantly lower levels of heterozygosity. Refuge populations also had considerably lower levels of allelic diversity than parental populations. All losses were of relatively rare alleles (frequency less than 0.1 in parental population). These losses were probably due to an undocumented bottleneck early in the introduction history. These results were surprising because the initial transplant involved 900 fish and because mosquitofish have numerous reproductive traits that should minimize the effects of bottlenecks on genetic diversity. A literature review revealed that genetic variability is often reduced in refuge populations and that such reductions typically involve the loss of alleles. We suggest that translocated populations be examined periodically for losses of genetic variability.     

Patterns of Genetic Diversity in Remaining Giant Panda Populations

Abstract: The giant panda ( Ailuropoda melanoleuca ) is among the more familiar symbols of species conservation. The protection of giant panda populations has been aided recently by the establishment of more and better-managed reserves in existing panda habitat located in six mountain ranges in western China. These remaining populations are becoming increasingly isolated from one another, however, leading to the concern that historic patterns of gene flow will be disrupted and that reduced population sizes will lead to diminished genetic variability. We analyzed four categories of molecular genetic markers (mtDNA restriction-fragment-length polymorphisms [RFLP], mtDNA control region sequences, nuclear multilocus DNA fingerprints, and microsatellite size variation) in giant pandas from three mountain populations (Qionglai, Minshan, and Qinling) to assess current levels of genetic diversity and to detect evidence of historic population subdivisions. The three populations had moderate levels of genetic diversity compared with similarly studied carnivores for all four gene measures, with a slight but consistent reduction in variability apparent in the smaller Qinling population. That population also showed significant differentiation consistent with its isolation since historic times. From a strictly genetic perspective, the giant panda species and the three populations look promising insofar as they have retained a large amount of genetic diversity in each population, although evidence of recent population reduction—likely from habitat loss—is apparent. Ecological management to increase habitat, population expansion, and gene flow would seem an effective strategy to stabilize the decline of this endangered species.     

Effective Population Size in Red-Cockaded Woodpeckers: Population and Model Differences

Loss of genetic variability in isolated populations is an important issue for conservation biology. Most studies involve only a single population of a given species and a single method of estimating rate of loss. Here we present analyses for three different Red-cockaded Woodpecker ( Picoides borealis ) populations from different geographic regions. We compare two different models for estimating the expected rate of loss of genetic variability, and test their sensitivity to model parameters. We found that the simpler model (Reed et al. 1988) consistently estimated a greater rate of loss of genetic variability from a population than did the Emigh and Pollak (1979) model. The ratio of effective population size (which describes the expected rate of loss of genetic variability) to breeder population size varied widely among Red-cockaded Woodpecker populations due to geographic variation in demography. For this species, estimates of effective size were extremely sensitive to survival parameters, but not to the probability of breeding or reproductive success. Sensitivity was sufficient that error in estimating survival rates in the field could easily mask true population differences in effective size. Our results indicate that accurate and precise demographic data are prerequisites to determining effective population size for this species using genetic models, and that a single estimate of rate of loss of genetic variability is not valid across populations.     

Erosion of Heterozygosity in Fluctuating Populations

Abstract: Demographic, environmental, and genetic stochasticity threaten the persistence of isolated populations. The relative importance of these intertwining factors remains unresolved, but a common view is that random demographic and environmental events will usually drive small populations to the brink of extinction before genetic deterioration poses a serious threat. To evaluate the potential importance of genetic factors, we analyzed a model linking demographic and environmental conditions to the loss of genetic diversity in isolated populations undergoing natural levels of fluctuation. Nongenetic processes—environmental stochasticity and population demography—were modeled according to a bounded diffusion process. Genetic processes were modeled by quantifying the rate of drift according to the effective population size, which was predicted from the same parameters used to describe the nongenetic processes. We combined these models to predict the heterozygosity remaining at the time of extinction, as predicted by the nongenetic portion of the model. Our model predicts that many populations will lose most or all of their neutral genetic diversity before nongenetic random events lead to extinction. Given the abundant evidence for inbreeding depression and recent evidence for elevated extinction rates of inbred populations, our findings suggest that inbreeding may be a greater general threat to population persistence than is generally recognized. Therefore, conservation biologists should not ignore the genetic component of extinction risk when assessing species endangerment and developing recovery plans.     

Conservation of Marine Megafauna through Minimization of Fisheries Bycatch

Abstract: Many populations of marine megafauna, including seabirds, sea turtles, marine mammals, and elasmobranchs, have declined in recent decades due largely to anthropogenic mortality. To successfully conserve these long‐lived animals, efforts must be prioritized according to feasibility and the degree to which they address threats with the highest relative impacts on population dynamics. Recently, Wilcox and Donlan (2007, Frontiers in Ecology and the Environment) and Donlan and Wilcox (2008, Biological Invasions) proposed a conservation strategy of “compensatory mitigation” in which fishing industries offset bycatch of seabirds and sea turtles by funding eradication of invasive mammalian predators from the terrestrial reproductive sites of these marine animals . Although this is a creative and conceptually compelling approach, we find it flawed as a conservation tool because it has narrow applicability among marine megafauna, it does not address the most pervasive threats to marine megafauna, and it is logistically and financially infeasible. Invasive predator eradication does not adequately offset the most pressing threat to most marine megafauna populations—fisheries bycatch. For seabird populations, fisheries bycatch and invasive predators infrequently are overlapping threats. Invasive predators have limited population‐level impacts on sea turtles and marine mammals and no impacts on elasmobranchs, all of which are threatened by bycatch. Implementing compensatory mitigation in marine fisheries is unrealistic due to inadequate monitoring, control, and surveillance in the majority of fleets. Therefore, offsetting fisheries bycatch with eradication of invasive predators would be less likely to reverse population declines than reducing bycatch. We recommend that efforts to mitigate bycatch in marine capture fisheries should address multiple threats to sensitive bycatch species groups, but these efforts should first institute proven bycatch avoidance and reduction methods before considering compensatory mitigation.     

Limited population structure in Northern and Spotted Wolffishes (<Emphasis Type="Italic">Anarhichas denticulatus</Emphasis> and <Emphasis Type="Italic">A. minor</Emphasis>) despite low apparent dispersal potential

Northern and Spotted Wolffishes (Anarhichas denticulatus and A. minor) are demersal marine fishes listed as “threatened” in Canadian waters. Both species have unusually large benthic eggs and large size at hatch, which should reduce passive dispersal. We examined population differentiation with microsatellite and AFLP loci across the ranges of both species in the North Atlantic Ocean. Although significant population structure was documented, differentiation was less than expected based on knowledge of life history characteristics. Significant differentiation was found in Northern Wolffish between the Barents Sea and other samples based on both microsatellite and AFLP data. In contrast, population structure in the Spotted Wolffish was notably weaker, particularly with microsatellites. Both species were characterized by low genetic diversity for marine fishes and had significantly lower genetic diversity than the congeneric Atlantic Wolffish. This finding was consistent with the conservation status of these three species and suggests potential vulnerability to over-exploitation in Northern and Spotted Wolffishes.     

Genetic population structure in the black-spot sea bream (<Emphasis Type="Italic">Pagellus bogaraveo</Emphasis> Brünnich, 1768) from the NE Atlantic

The depletion of shallow-water fish stocks through overexploitation has led to increasing fishing pressure on deep-sea species. Poor knowledge of the biology of commercially valuable deep-water fish has led to the serial depletion of stocks of several species across the world. Data regarding the genetic structure of deep-sea fish populations is important in determining the impact of overfishing on the overall genetic variability of species and can be used to estimate the likelihood of recolonisation of damaged populations through immigration of individuals from distant localities. Here the genetic structure of the commercially fished deep-water species the blackspot sea bream, Pagellus bogaraveo is investigated in the northeastern Atlantic using partial DNA sequencing of mitochondrial cytochrome b (cyt-b) and D-loop regions and genotyping of microsatellite loci. An absence of variation in cyt-b and low genetic variation in D-loop sequences potentially indicate that P. bogaraveo may have undergone a severe bottleneck in the past. Similar bottlenecks have been detected in other Atlantic species of fish and have possibly originated from the last glaciation. P. bogaraveo may have been particularly vulnerable to the effects of low temperature and a fall in sea level because stages of its life history occur in shallow water and coastal sites. However, there are other explanations of low genetic variability in populations of P. bogaraveo, such as a low population size and the impacts of fishing on population structure. Analysis of population structure using both D-loop and microsatellite analysis indicates low to moderate, but significant, genetic differentiation between populations at a regional level. This study supports studies on other deep-sea fish species that indicate that hydrographic or topographic barriers prevent dispersal of adults and/or larvae between populations at regional and oceanographic scales. The implications for the management and conservation of deep-sea fish populations are discussed.Communicated by J.P. Thorpe, Port Erin     

Genetic and demographic consequences of importing animals into a small population: a simulation model of the Texas State Bison Herd (USA)

The extant 40 bison (Bison bison) constituting the Texas State Bison Herd (TSBH; USA) are directly and exclusively descended from a bison herd assembled by Charles Goodnight in the 1880s, representing a historically and genetically valuable resource. The population currently suffers from low genetic variation, low heterozygosity, high calf mortality, and low natality rates compared with other closed bison populations. Population viability analysis using the VORTEX program previously indicated a 99% chance of population extinction within the next 41 years [J. Mamm. 85 (2004) in press]. We developed a stochastic simulation model to evaluate the genetic and demographic consequences of various management scenarios for the TSBH using genotypic data from 51 microsatellite loci and demographic information recorded over a 6-year period. Our results reveal that without the introduction of new genetic variation, approximately 37% of the representative microsatellite loci will become fixed as the TSBH continues to lose genetic variation at a staggering rate of 30–40% within the next 50 years. Furthermore, if the current trends in natality and mortality rates continue, our model indicates the TSBH faces a 99% chance of extinction in the next 51 years. With the importation of unrelated male bison into the TSBH, and under the assumption of increased fitness, the probability of population survival in the next 100 years increases to 100%, and the population will reach the approximate carrying capacity of 200 bison in 15–16 years. Furthermore, our model predicts increases in genetic diversity and heterozygosity of 24.7–48.4% and 17.5–36.5%, respectively, in the next 100 years following the addition of new genetic variation. We conclude that the importation of bison into the TSBH is necessary to prevent extinction and ensure long-term population survival.     

Human Impacts on Drainages of the Mesa Central, Mexico, and Its Genetic Effects on an Endangered Fish, Zoogoneticus quitzeoensis

Abstract:  The Mesa Central of Mexico is of special conservation interest due to its high richness of freshwater fish species, of which the goodeines are one of the most representative groups. Through an integrated approach, we determined conservation priorities for goodeine populations. We based our recommendations on the genetic diversity (variation in five microsatellite DNA loci) in 10 populations of Zoogoneticus quitzeoensis and on an analysis of ecological (e.g., presence of exotic species), social (e.g., political situation), and environmental (e.g., pollution) information for 52 historical occurrence points for species in the genus Zoogoneticus . Patterns of genetic erosion and genetic diversity indices were closely associated with human impact. Recent bottleneck events were most evident in the populations from remnants of the lakes drained at the beginning of the twentieth century. We identified seven operational conservation units (OCUs), all of which should be conserved because they contain unique portions of the total variation of the species. Special attention needs to be given to increase genetic variability, recover population sizes, and reestablish contact among populations within OCUs. It is imperative to create an integrative and effective approach for the recovery and conservation of the freshwater fish diversity of Central Mexico that is based on social and natural sciences.     

Conservation Implications of Drastic Reductions in the Smallest and Most Isolated Populations of Giant Pandas

Abstract: In conservation biology, understanding the causes of endangerment is a key step to devising effective conservation strategies. We used molecular evidence (coalescent simulations of population changes from microsatellite data) and historical information (habitat and human population changes) to investigate how the most‐isolated populations of giant pandas (Ailuropoda melanoleuca) in the Xiaoxiangling Mountains became highly endangered. These populations experienced a strong, recent demographic reduction (60‐fold), starting approximately 250 years BP. Explosion of the human population and use of non‐native crop species at the peak of the Qing Empire resulted in land‐use changes, deforestation, and habitat fragmentation, which are likely to have led to the drastic reduction of the most‐isolated populations of giant pandas. We predict that demographic, genetic, and environmental factors will lead to extinction of giant pandas in the Xiaoxiangling Mountains in the future if the population remains isolated. Therefore, a targeted conservation action—translocation—has been proposed and is being implemented by the Chinese goverment.     

Environmental Surrogates for Predicting and Conserving Adaptive Genetic Variability in Tree Species

Abstract: Adaptive genetic variability within species is an essential component of biodiversity but has been largely ignored in studies aimed at assessing and predicting biodiversity of the forest environment. We used factorial regression and structuring models to test easily measured surrogates, such as ecological attributes, as predictors of adaptive genetic variation between populations of a tree species ( Eucalyptus delegatensis ). Adaptive variability was defined in terms of variation in average growth performance of 68 populations and of population-by-environment interaction across seven different environments. The best surrogates of genetic variability were measures of solar radiation and temperature range, each predicting more than 50% of the genetic variability within the species. Rock and understory types, when used either alone or in combination with other covariates, also were very efficient in discriminating between populations in groups showing similar adaptation. Significant relationships between particular surrogates and growth patterns of variation were attributed to effects of natural selection that had occurred in the population source locations. We recommend the development of studies focusing on the population level of biodiversity to improve the conservation of forest ecosystems in Australia.     

Contrasting population structures in two sympatric fishes in the Baltic Sea basin

Detailed multispecies studies on the patterns of genetic variability and differentiation in marine environments are still rare. Using mitochondrial and nuclear genetic markers, we compared genetic variability and population structuring of threespine (Gasterosteus aculeatus) and ninespine (Pungitius pungitius) sticklebacks from the same eleven marine and six freshwater locations within the Baltic Sea basin. Analyses of both marker types revealed a significantly lower degree of genetic structuring in both marine and freshwater populations of threespine than those ninespine sticklebacks. Isolation-by-distance (IBD) was detected across the marine populations in both species, suggesting spatially limited gene flow. However, the levels of genetic diversity and differentiation across the localities were uncorrelated between the two species in both marine and freshwater environments. Accordingly, estimates of effective population sizes were larger and migration rates were higher for three- than for ninespine sticklebacks. Hence, ninespine stickleback populations from the Baltic Sea basin appear to be subject to stronger genetic drift than sympatric threespine sticklebacks, and the proximate reason for this difference is likely to be found from autecological differences between the two species. In accordance with the earlier studies, genetic variability was higher and the degree of genetic differentiation was lower in marine than in freshwater populations in both species.     

Relationship of Genetic Variation to Population Size in Wildlife

Genetic diversity is one of three levels of biological diversity requiring conservation. Genetic theory predicts that levels of genetic variation should increase with effective population size. Soulé (1976) compiled the first convincing evidence that levels of genetic variation in wildlife were related to population size, but this issue remains controversial. The hypothesis that genetic variation is related to population size leads to the following predictions: (1) genetic variation within species should be related to population size; (2) genetic variation within species should be related to island size; (3) genetic variation should be related to population size within taxonomic groups; (4) widespread species should have more genetic variation than restricted species; (5) genetic variation in animals should be negatively correlated with body size; (6) genetic variation should be negatively correlated with rate of chromosome evolution; (7) genetic variation across species should be related to population size; (8) vertebrates should have less genetic variation than invertebrates or plants; (9) island populations should have less genetic variation than mainland populations; and (10) endangered species should have less genetic variation than nonendangered species. Empirical observations support all these hypotheses. There can be no doubt that genetic variation is related to population size, as Soulé proposed. Small population size reduces the evolutionary potential of wildlife species.     

Genetic diversity and IUCN Red List status

The International Union for Conservation of Nature (IUCN) Red List is an important and widely used tool for conservation assessment. The IUCN uses information about a species’ range, population size, habitat quality and fragmentation levels, and trends in abundance to assess extinction risk. Genetic diversity is not considered, although it affects extinction risk. Declining populations are more strongly affected by genetic drift and higher rates of inbreeding, which can reduce the efficiency of selection, lead to fitness declines, and hinder species’ capacities to adapt to environmental change. Given the importance of conserving genetic diversity, attempts have been made to find relationships between red-list status and genetic diversity. Yet, there is still no consensus on whether genetic diversity is captured by the current IUCN Red List categories in a way that is informative for conservation. To assess the predictive power of correlations between genetic diversity and IUCN Red List status in vertebrates, we synthesized previous work and reanalyzed data sets based on 3 types of genetic data: mitochondrial DNA, microsatellites, and whole genomes. Consistent with previous work, species with higher extinction risk status tended to have lower genetic diversity for all marker types, but these relationships were weak and varied across taxa. Regardless of marker type, genetic diversity did not accurately identify threatened species for any taxonomic group. Our results indicate that red-list status is not a useful metric for informing species-specific decisions about the protection of genetic diversity and that genetic data cannot be used to identify threat status in the absence of demographic data. Thus, there is a need to develop and assess metrics specifically designed to assess genetic diversity and inform conservation policy, including policies recently adopted by the UN's Convention on Biological Diversity Kunming-Montreal Global Biodiversity Framework.     

Spatial heterogeneity in distribution and ecology of Western Palearctic birds

Species vary in abundance and heterogeneity of spatial distribution, and the ecological and evolutionary consequences of such variability are poorly known. Evolutionary adaptation to heterogeneously distributed resources may arise from local adaptation with individuals of such locally adapted populations rarely dispersing long distances and hence having small populations and small overall ranges. We quantified mean population density and spatial heterogeneity in population density of 197 bird species across 12 similarly sized regions in the Western Palearctic. Variance in population density among regions differed significantly from a Poisson distribution, suggesting that random processes cannot explain the observed patterns. National estimates of means and variances in population density were positively correlated with continental estimates, suggesting that means and variances were maintained across spatial scales. We used Morisita's index of population abundance as an estimate of heterogeneity in distribution among regions to test a number of predictions. Heterogeneously distributed passerine bird species as reflected by Morisita's index had small populations, low population densities, and small breeding ranges. Their breeding populations had been consistently maintained at low levels for considerable periods of time, because the degree of genetic variation in a subsample of non-passerines and passerines was significantly negatively related to heterogeneity in distribution. Heterogeneously distributed passerine species were not more often habitat specialists than homogeneously distributed species. Furthermore, heterogeneously distributed passerine species had high annual adult survival rates but did not differ in annual fecundity from homogeneously distributed species. Heterogeneously distributed passerine species rarely colonized urban habitats. Finally, homogeneously distributed bird species were hosts to a greater diversity of blood parasite species than heterogeneously distributed species. In conclusion, small breeding ranges, population sizes, and population densities of heterogeneously distributed passerine bird species, combined with their low degree of genetic variability, and their inability to colonize urban areas may render such species particularly susceptible to human-influenced global climatic changes.     

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.