Genetic differentiation among populations of a broadcast spawning soft coral, Sinularia flexibilis, on the Great Barrier Reef

The genetic structure of 12 reef populations of the soft coral Sinularia flexibilis (Octocorallia, Alcyoniidae) was studied along the Great Barrier Reef (GBR) at a maximum separation of 1,300 km to investigate the relative importance of sexual and asexual reproduction, genetic differentiation and gene flow among these populations. S. flexibilis is a widely distributed Indo-Pacific species and a gamete broadcaster that can form large aggregations of colonies on near-shore reefs of the GBR. Up to 60 individuals per reef were collected at a minimum sampling scale of 5 m at two sites per reef, from December 1998 to February 2000. Electrophoretic analyses of nine polymorphic allozymes indicated that genotypic frequencies in most populations and loci did not differ significantly from those expected from Hardy–Weinberg predictions. Analysis of multi-locus genotypes indicated a high number of unique genotypes (N _go) relative to the number of individuals sampled (N) in each reef population (range of 0.69–0.95). The maximum number of individuals likely to have been produced sexually (N*) was similar to the number of individuals sampled (i.e. N*:N ˜ 1), suggesting that even repeated genotypes may have been produced sexually. These results demonstrated a dominant role of sexual reproduction in these populations at the scale sampled. Significant genetic differentiation between some populations indicated that gene flow is restricted between some reefs (F _ST=0.026, 95% CI= 0.011 − 0.045) and even between sites within reefs (F _ST=0.041, 95% CI=0.027 − 0.055). Nevertheless, there was no relationship between geographic separation and genetic differentiation. Analyses comparing groups of populations showed no significant differentiation on a north-south gradient in the GBR. The pattern in the number of significant differences in gene frequencies in pairwise population comparisons, however, suggested that gene flow may be more restricted among inner-shelf reef populations near to the coast than among mid/outer-shelf populations further from the coast. Received: 10 July 2000 / Accepted: 5 October 2000     

Range‐Wide Genetic Connectivity of the Hawaiian Monk Seal and Implications for Translocation

Abstract: The Hawaiian monk seal (Monachus schauinslandi) is one of the most critically endangered marine mammals. Less than 1200 individuals remain, and the species is declining at a rate of approximately 4% per year as a result of juvenile starvation, shark predation, and entanglement in marine debris. Some of these problems may be alleviated by translocation; however, if island breeding aggregates are effectively isolated subpopulations, moving individuals may disrupt local adaptations. In these circumstances, managers must balance the pragmatic need of increasing survival with theoretical concerns about genetic viability. To assess range‐wide population structure of the Hawaiian monk seal, we examined an unprecedented, near‐complete genetic inventory of the species (n =1897 seals, sampled over 14 years) at 18 microsatellite loci. Genetic variation was not spatially partitioned (_w=?0.03, p = 1.0), and a Bayesian clustering method provided evidence of one panmictic population (K =1). Pairwise F_ST comparisons (among 7 island aggregates over 14 annual cohorts) did not reveal temporally stable, spatial reproductive isolation. Our results coupled with long‐term tag‐resight data confirm seal movement and gene flow throughout the Hawaiian Archipelago. Thus, human‐mediated translocation of seals among locations is not likely to result in genetic incompatibilities.     

Hunting Effects on Favourable Conservation Status of Highly Inbred Swedish Wolves

The wolf (Canis lupus) is classified as endangered in Sweden by the Swedish Species Information Centre, which is the official authority for threat classification. The present population, which was founded in the early 1980s, descends from 5 individuals. It is isolated and highly inbred, and on average individuals are more related than siblings. Hunts have been used by Swedish authorities during 2010 and 2011 to reduce the population size to its upper tolerable level of 210 wolves. European Union (EU) biodiversity legislation requires all member states to promote a concept called “favourable conservation status” (FCS) for a series of species including the wolf. Swedish national policy stipulates maintenance of viable populations with sufficient levels of genetic variation of all naturally occurring species. Hunting to reduce wolf numbers in Sweden is currently not in line with national and EU policy agreements and will make genetically based FCS criteria less achievable for this species. We suggest that to reach FCS for the wolf in Sweden the following criteria need to be met: (1) a well‐connected, large, subdivided wolf population over Scandinavia, Finland, and the Russian Karelia‐Kola region should be reestablished, (2) genetically effective size (N_e) of this population is in the minimum range of N_e = 500–1000, (3) Sweden harbors a part of this total population that substantially contributes to the total N_e and that is large enough to not be classified as threatened genetically or according to IUCN criteria, and (4) average inbreeding levels in the Swedish population are <0.1. Efectos de la Cacería sobre el Estatus de Conservación Favorable de Lobos Suecos con Endogamia Alta     

Phylogeography and Limited Genetic Connectivity in the Endangered Boring Giant Clam across the Coral Triangle

Abstract: The Coral Triangle is the global center of marine biodiversity; however, its coral reefs are critically threatened. Because of the bipartite life history of many marine species with sedentary adults and dispersive pelagic larvae, designing effective marine protected areas requires an understanding of patterns of larval dispersal and connectivity among geographically discrete populations. We used mtDNA sequence data to examine patterns of genetic connectivity in the boring giant clam (Tridacna crocea) in an effort to guide conservation efforts within the Coral Triangle. We collected an approximately 485 base pair fragment of mtDNA cytochrome c oxidase 1 (CO1) from 414 individuals at 26 sites across Indonesia. Genetic structure was strong between regions (φ_ST=0.549, p < 0.00001) with 3 strongly supported clades: one restricted to western Sumatra, another distributed across central Indonesia, and a third limited to eastern Indonesia and Papua. Even within the single largest clade, small but significant genetic structure was documented (φ_ST=0.069, p < 0.00001), which indicates limited gene flow within and among phylogeographic regions. Significant patterns of isolation by distance indicated an average dispersal distance of only 25–50 km, which is far below dispersal predictions of 406–708 km derived from estimates of passive dispersal over 10 days via surface currents. The strong regional genetic structure we found indicates potent limits to genetic and demographic connectivity for this species throughout the Coral Triangle and provides a regional context for conservation planning. The recovery of 3 distinct evolutionarily significant units within a well‐studied taxonomic group suggests that biodiversity in this region may be significantly underestimated and that Tridacna taxa may be more endangered than currently recognized.     

Restricted gene flow and evolutionary divergence between geographically separated populations of the Antarctic octopus Pareledone turqueti

Samples of the Antarctic octopus Pareledone turqueti were taken from three locations on the Scotia Ridge in the Southern Ocean. The genetic homogeneity of these populations was investigated using isozyme electrophoresis. Whilst panmixia appeared to be maintained around South Georgia (F _ST = 0) gene flow between this island and Shag Rocks, an island only 150 km away but separated by great depths, was extremely limited (F _ST = 0.74). These results are examined with respect to the discontinuous distribution of P. turqueti throughout Antarctica. An estimate of effective population size was also calculated (N _e = 3600). Received: 7 March 1997 / Accepted: 27 March 1997     

Use of Habitats as Surrogates of Biodiversity for Efficient Coral Reef Conservation Planning in Pacific Ocean Islands

Abstract: Marine protected areas (MPAs) have been highlighted as a means toward effective conservation of coral reefs. New strategies are required to more effectively select MPA locations and increase the pace of their implementation. Many criteria exist to design MPA networks, but generally, it is recommended that networks conserve a diversity of species selected for, among other attributes, their representativeness, rarity, or endemicity. Because knowledge of species’ spatial distribution remains scarce, efficient surrogates are urgently needed. We used five different levels of habitat maps and six spatial scales of analysis to identify under which circumstances habitat data used to design MPA networks for Wallis Island provided better representation of species than random choice alone. Protected‐area site selections were derived from a rarity–complementarity algorithm. Habitat surrogacy was tested for commercial fish species, all fish species, commercially harvested invertebrates, corals, and algae species. Efficiency of habitat surrogacy varied by species group, type of habitat map, and spatial scale of analysis. Maps with the highest habitat thematic complexity provided better surrogates than simpler maps and were more robust to changes in spatial scales. Surrogates were most efficient for commercial fishes, corals, and algae but not for commercial invertebrates. Conversely, other measurements of species‐habitat associations, such as richness congruence and composition similarities provided weak results. We provide, in part, a habitat‐mapping methodology for designation of MPAs for Pacific Ocean islands that are characterized by habitat zonations similar to Wallis. Given the increasing availability and affordability of space‐borne imagery to map habitats, our approach could appreciably facilitate and improve current approaches to coral reef conservation and enhance MPA implementation.     

Understanding and Estimating Effective Population Size for Practical Application in Marine Species Management

Abstract: Effective population size (N_e) determines the strength of genetic drift in a population and has long been recognized as an important parameter for evaluating conservation status and threats to genetic health of populations. Specifically, an estimate of N_e is crucial to management because it integrates genetic effects with the life history of the species, allowing for predictions of a population's current and future viability. Nevertheless, compared with ecological and demographic parameters, N_e has had limited influence on species management, beyond its application in very small populations. Recent developments have substantially improved N_e estimation; however, some obstacles remain for the practical application of N_e estimates. For example, the need to define the spatial and temporal scale of measurement makes the concept complex and sometimes difficult to interpret. We reviewed approaches to estimation of N_e over both long‐term and contemporary time frames, clarifying their interpretations with respect to local populations and the global metapopulation. We describe multiple experimental factors affecting robustness of contemporary N_e estimates and suggest that different sampling designs can be combined to compare largely independent measures of N_e for improved confidence in the result. Large populations with moderate gene flow pose the greatest challenges to robust estimation of contemporary N_e and require careful consideration of sampling and analysis to minimize estimator bias. We emphasize the practical utility of estimating N_e by highlighting its relevance to the adaptive potential of a population and describing applications in management of marine populations, where the focus is not always on critically endangered populations. Two cases discussed include the mechanisms generating N_e estimates many orders of magnitude lower than census N in harvested marine fishes and the predicted reduction in N_e from hatchery‐based population supplementation.     

Expanding marine protected areas to include degraded coral reefs

Marine protected areas (MPAs) are a commonly applied solution to coral reef degradation, yet coral reefs continue to decline worldwide. We argue that expanding the range of MPAs to include degraded reefs (DR‐MPA) could help reverse this trend. This approach requires new ecological criteria for MPA design, siting, and management. Rather than focusing solely on preserving healthy reefs, our approach focuses on the potential for biodiversity recovery and renewal of ecosystem services. The new criteria would help identify sites with the highest potential for recovery and the greatest resistance to future threats (e.g., increased temperature and acidification) and sites that contribute to MPA connectivity. The DR‐MPA approach is a compliment rather than a substitute for traditional MPA design approaches. We believe that the DR‐MPA approach can enhance the natural, or restoration‐assisted, recovery of DRs and their ecosystem services; increase total reef area available for protection; promote more resilient and better‐connected MPA networks; and improve conditions for human communities dependent on MPA ecosystem services.     

Population genetic structure and genetic diversity of three critically endangered <Emphasis Type="Italic">Pristis</Emphasis> sawfishes in Australian waters

Northern Australia is considered to be one of the last strongholds for three critically endangered sawfishes, Pristis zijsron, Pristis clavata, and Pristis microdon, making these populations of global significance. Population structure and levels of genetic diversity were assessed for each species across northern Australia using a portion of the mitochondrial control region. Statistically significant genetic structure was detected in all three species, although it was higher in P. microdon (F _ST = 0.811; N = 149) than in either P. clavata (F _ST = 0.419; N = 73) or P. zijsron (F _ST = 0.202; N = 49), possibly due to a much higher and/or localized level of female philopatry in P. microdon. The overall levels of haplotype diversity in P. zijsron (h = 0.555), P. clavata (h = 0.489), and P. microdon (h = 0.650) were moderate, although it appears to be reduced in the assemblages of P. zijsron and P. clavata in the Gulf of Carpentaria (h = 0.342 and h = 0.083, respectively). Since female migration (replenishment) between regions is unlikely, conservation plans should strive to maintain current levels of diversity and abundances in the regional assemblages of each species.     

Local and regional genetic connectivity in a Caribbean coral reef fish

Coupled bio-physical models of larval dispersal predict that the Costa Rica–Panama (CR–PAN) reefs should constitute a demographically isolated region in the western Caribbean. We tested the hypothesis that CR–PAN coral reef fish populations would be isolated from Mesoamerican Barrier Reef System (MBRS) populations. To test that, we assessed population genetic structure in bicolor damselfish (Stegastes partitus) from both regions. Adult fish were genotyped from five reefs in CR–PAN and from four reefs along the MBRS at 12 microsatellite loci. Between-region F _ST (F _ST = 0.0030, P < 0.005) and exact test (x ² = 74.34, df = 18, P < 0.0001) results indicated that there is weak but significant genetic differentiation between regions, suggesting some restriction in connectivity along the Central American coastline, as predicted by bio-oceanographic models. Additionally, there is among-site genetic structure in the CR–PAN region, relative to the MBRS and between regions, suggesting higher self-recruitment within CR–PAN. This finding may be explained by differences in habitat characteristics.     

Mode of reproduction,recruitment, and genetic subdivision in the brooding sponge<Emphasis Type="Italic"> Haliclona</Emphasis> sp.

Sponges display a variety of reproductive strategies that have the potential to influence population genetic structure. Histological examination of ten reproductive individuals of the Western Australian sponge Haliclona sp. showed that this species broods embryonic larvae that are potentially limited in dispersal capabilities. Because sponges have the potential to propagate in a number of modes, allozyme electrophoresis was used to assess the relative importance of asexual and sexual reproduction to recruitment, and to quantify genetic subdivision over different spatial scales. Tissue samples from 227 sponges were collected from reefs within two areas 400 km apart: Hamelin Bay and Rottnest Island. Contrary to expectations for highly clonal populations, genotypic diversity within sites was high, no linkage disequilibrium was found, and there was no evidence of genotypic clustering within reefs. There was no genetic evidence that asexual reproduction is important for the maintenance of populations. Genetic comparisons were consistent with mixing of sexually produced recruits within reefs, on a scale up to a few hundred metres, but significant genetic subdivision between reefs (F_ST=0.069 at Hamelin Bay, 0.130 at Rottnest Island) indicated that water gaps of several hundred metres are effective at preventing dispersal. Subdivision between the two areas, separated by 400 km, was moderately greater (F_ST=0.142) than within, but the same alleles were predominant in the two areas. These genetic patterns are consistent with limited dispersal capabilities of brooded larvae.Communicated by G.F. Humphrey, Sydney     

Genetic differentiation of bicolor damselfish (Eupomacentrus partitus) populations in the Florida Keys

Electrophoretic variation in proteins encoded by 23 loci revealed substantial genetic differentiation among populations of bicolor damselfish (Eupomacentrus partitus) collected from four coral reefs in the Florida Keys, USA, during 1986–1988. Genetic differentiation was concentrated between a sample collected from Little Grecian Rocks Reef (LGR) and the remaining samples, including fish from a reef only 600 m distant (Grecian Rocks Reef). Genetic distinction of the LGR sample derived from significantly heterogenous allelic frequencies at six of eight polymorphic loci. Aco-1 (aconitase); Ada (adenosine deaminase); Gpi-2 (glucosephosphate isomerase); Ldh-2 and Ldh-3 (lactate dehydrogenase); and Me-1 (malic enzyme); nevertheless, differentiation at cytosolic aconitase (Aco-1) far exceeded that observed for other loci (fixation index, F _ST=0.482), and differences in Aco-1 allele frequencies were largely responsible for large genetic distances (0.20) between LGR and the other reefs. Paradoxically, estimates of numbers of migrants exchanged between reefs per generation (mN _e=17.47) indicated the potential for extensive gene flow. The extent of genetic differentiation among these populations is evaluated relative to models of population genetic structure based on equilibrium between gene flow and natural selection or genetic drift.     

Population genetics of Trochus niloticus and Tectus coerulescens,topshells with short-lived larvae

Trochus niloticus L. and Tectus coerulescens Lmk., two coral reef trochid gastropods that have similar life-histories including a lecithotrophic larval stage, were sampled from reefs in the northern, central and southern sections of the Great Barrier Reef (GBR) in 1991. Significant sex-ratio biases were noted, and these varied among reefs, apparently with latitude. Demographic data suggested that highly discontinuous and localized recruitment occurs. Surveys of allozyme frequencies at 12 loci revealed no significant genetic differences among populations of Trochus niloticus at any geographical scale. High gene flow between zones of the GBR was inferred, with the number of migrants per generation (N _m ) of the order of 100. Directional selection was thought to occur at one locus (GDH ^*). Genetic variability in T. niloticus (H=0.069 to 0.110, and only three loci polymorphic) was low compared with other trochids. This was thought to be due to smaller effective population size, resulting from an unbalanced sexratio, aggregative spatial distribution of adults, high variance in reproductive success, and/or the occurrence of population extinctions and recolonisations. In contrast, Tectus coerulescens exhibited significant genetic differences between zones, indicating a much lower rate of migration between populations (N _m 1 to 10), and displayed high genetic diversity (H=0.225 to 0.279). A trend for increasing genetic diversity from the northern to southern GBR was found in both species. It is not clear whether the occurrence of two contrasted population genetic structures in species with apparently similar life-histories is due to ecological or historical factors.     

Spatial,socio‐economic,and ecological implications of incorporating minimum size constraints in marine protected area network design

Marine protected areas (MPAs) are the cornerstone of most marine conservation strategies, but the effectiveness of each one partly depends on its size and distance to other MPAs in a network. Despite this, current recommendations on ideal MPA size and spacing vary widely, and data are lacking on how these constraints might influence the overall spatial characteristics, socio‐economic impacts, and connectivity of the resultant MPA networks. To address this problem, we tested the impact of applying different MPA size constraints in English waters. We used the Marxan spatial prioritization software to identify a network of MPAs that met conservation feature targets, whilst minimizing impacts on fisheries; modified the Marxan outputs with the MinPatch software to ensure each MPA met a minimum size; and used existing data on the dispersal distances of a range of species found in English waters to investigate the likely impacts of such spatial constraints on the region's biodiversity. Increasing MPA size had little effect on total network area or the location of priority areas, but as MPA size increased, fishing opportunity cost to stakeholders increased. In addition, as MPA size increased, the number of closely connected sets of MPAs in networks and the average distance between neighboring MPAs decreased, which consequently increased the proportion of the planning region that was isolated from all MPAs. These results suggest networks containing large MPAs would be more viable for the majority of the region's species that have small dispersal distances, but dispersal between MPA sets and spill‐over of individuals into unprotected areas would be reduced. These findings highlight the importance of testing the impact of applying different MPA size constraints because there are clear trade‐offs that result from the interaction of size, number, and distribution of MPAs in a network.     

Genetic consequences of long larval life in the starfish Linckia laevigata (Echinodermata: Asteroidea) on the Great Barrier Reef

Gene flow between populations of the asteroid Linckia laevigata (Linnaeus) was investigated by examining over 1000 individuals collected from ten reefs throughout the Great Barrier Reef (GBR), Australia, for genetic variation at seven polymorphic enzyme loci. Despite geographic separations in excess of 1000 km, Nei's unbiased genetic distance (0 to 0.003) and standardised genetic variation between populations (F _ST) values (mean 0.0011) were small and not significant. Genetic homogeneity among L. laevigata populations is consistent with the long-distance dispersal capability of its 28 d planktonic larval phase, and is greater than that observed for other asteroid species, including another high-dispersal species, Acanthaster planci, which has a 14 d larval phase. Variation within populations was also higher than previously recorded for asteroids (mean heterozygosity=0.384; number of alleles per locus ranged from 5.1 to 6.0 in each population). Among asteroids, dispersal ability is positively correlated with gene flow and levels of variation, and negatively correlated with levels of differentiation.     

Population genetics of the nurse shark (Ginglymostoma cirratum) in the western Atlantic

The nurse shark, Ginglymostoma cirratum, inhabits shallow, tropical, and subtropical waters in the Atlantic and the eastern Pacific. Unlike many other species of sharks, nurse sharks are remarkably sedentary. We assayed the mitochondrial control region and eight microsatellite loci from individuals collected primarily in the western Atlantic to estimate the degree of population subdivision. Two individuals from the eastern Atlantic and one from the Pacific coast of Panama also were genotyped. Overall, the mtDNA haplotype (h = 48 ± 5%) and nucleotide (π = 0.08 ± 0.06%) diversities were low. The microsatellite data mirror the mitochondrial results with the average number of alleles ([`(N)]<sub>A</sub> \bar{N}_{A}  = 9) and observed heterozygosity ([`(H)]_O \bar{H}_{O}  = 0.58) both low. The low levels of diversity seen in both the mtDNA and the microsatellite may be due to historical sea level fluctuations and concomitant loss of shallow water habitat. Eight of the 10 pair-wise western Atlantic F _ST estimates for mtDNA indicated significant genetic subdivision. Pair-wise F _ST values for the microsatellite loci indicated a similar pattern as the mtDNA. The western Atlantic population of nurse sharks is genetically subdivided with the strongest separation seen between the offshore islands and mainland Brazil, likely due to deep water acting as a barrier to dispersal. The eastern and western Atlantic populations were closely related. The eastern Pacific individual is quite different from Atlantic individuals and may be a cryptic, sister species.     

Genetic variation and clonal structure in the scleractinian coral Pocillopora damicornis in the Ryukyu Archipelago, southern Japan

Samples of the scleractinian coral Pocillopora damicornis were collected from six sites located around four islands in the Ryukyu Archipelago, southern Japan, and subjected to allozyme electrophoresis. Seven polymorphic loci were examined for their allelic patterns. The ratio of observed to expected genotypic diversity (0.30 < G _o :G _e  < 0.64), the ratio of the observed number of genotypes to the number of individuals (0.47 < N _g :N _i  < 0.75), and deviations from Hardy–Weinberg equilibrium indicated that asexual reproduction plays a major role in the maintenance of established populations. However, populations were not completely dominated by a single or a few clones, and most clones were represented by only a few individual samples. The high frequency of typhoons in the region suggests that, in P. damicornis, fragmentation caused through occasional exposure to powerful waves is a major mode of asexual reproduction, but asexual production of planulae may also be contributing to the maintenance of populations. A significant genetic differentiation (F ST) was found between the six populations examined (0.027 < F ST < 0.092, average F ST = 0.056). The moderate gene flow is discussed according to characteristics of the larval stage of the species, and to circulation patterns in the region. Received: 7 August 1998 / Accepted: 18 May 1999     

Limited gene flow in the brooding coral Favia fragum (Esper, 1797)

Understanding population connectivity in corals is particularly important as these organisms are increasingly threatened by abiotic and biotic factors. This study examined the population genetic structure of the brooding coral Favia fragum across four locations in the Caribbean and Western Atlantic using mitochondrial and nuclear markers. Morphological features were also compared to test whether genetic diversity corresponds with skeletal morphology. When comparing across distantly related Caribbean and Bermudian locations, F _ST values were high and significant, indicating strong genetic structure. At a local scale, significant genetic structure was found among reefs in Panama, while no genetic structure was found among reefs within Barbados, Bermuda or Jamaica. Surprisingly, a single haplotype for each of the three markers examined was found in Bermuda, where samples varied significantly from all other locations in three out of four morphological features analyzed. These data indicate that gene flow of F. fragum may occur locally among reefs but is highly restricted at distant locations. Furthermore, isolated populations, such as that of Bermuda, must be self-seeding to maintain the observed genetic uniformity.     

Population structure and genetic variation in two tropical sea anemones (Cnidaria,Actinidae) with different reproductive strategies

Genetic variation and population structure of two tropical sea anemones, Bunodosoma caissarum Correa and Actinia bermudensis McMurrich were related to their different dispersal capabilities and reproductive modes. B. caissarum reproduces sexually and has a long-lived planktotrophic larva; A. bermudensis can reproduce both sexually and asexually, supposedly with short-range dispersal. Both species were sampled along 1150 km of Brazilian coastline between 1990 and 1991 and analyzed by horizontal starch gel electrophoresis for 16 enzyme loci in B. caissarum and 19 in A. bermudensis. B. caissarum had higher levels of heterozygosity (H=0.35) and lower levels of population structuring (F _ST=0.042) than A. bermudensis (H=0.17; F _ST=0.262). At one locality large genetic differences between two sympatric colour morphs of A. bermudensis provide evidence of possible cryptic speciation. Overall levels of genetic variation and heterozygosity in the two species are compatible with their known modes of reproduction.     

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.