Mitochondrial DNA Variability in Italian and East European Wolves: Detecting the Consequences of Small Population Size and Hybridization

Abstract: The Italian wolf ( Canis lupus ) population has declined continuously over the last few centuries and become isolated as a result of the extermination of other populations in central Europe and the Alps during the nineteenth century. In the 1970s, approximately 100 wolves survived in 10 isolated areas in the central and southern Italian Apennines. Loss of genetic variability, as suggested by preliminary studies of mitochondrial DNA (mtDNA) sequences, hybridization with feral dogs, and the illegal release of captive, non-native wolves are considered potential threats to the viability of the Italian wolf population. We sequenced 546 base pairs of the mtDNA control region in a comprehensive set of Italian wolves and compared them to those of dogs and other wolf populations from Europe and the Near East. Our data confirm the absence of mtDNA variability in Italian wolves: all 101 individuals sampled across their distribution in Italy had the same, unique haplotype, whereas seven haplotypes were found in only 26 wolves from an outbred population in Bulgaria. Most haplotypes were specific either to wolves or dogs, but some east European wolves shared haplotypes with dogs, indicative of hybridization. In contrast, neither hybridization with dogs nor introgression of non-native wolves was detected in the Italian population. These findings exclude the introgression of dog genes via matings between male wolves and female dogs, the most likely direction of hybridization. The observed mtDNA monomorphism is the possible outcome of random drift in the declining and isolated Italian wolf population, which probably existed at low effective population size during the last 100–150 years. Low effective population size and the continued loss of genetic variability might be a major threat to the long-term viability of Italian wolves. A controlled demographic increase, leading to recolonization of the historical wolf range in Italy, should be enforced.     

Genetic Variation of Naturally Colonizing Wolves in the Central Rocky Mountains

Recovery of gray wolf ( Canis lupus ) populations in North America depends on minimizing human-caused mortality and enhancing migration from stable source populations to suitable habitat unoccupied by wolves. We used a combination of field observation and DNA microsatellite genotyping to examine natural wolf colonization of Glacier National Park, Montana, and surrounding lands. We found high genetic variation in the colonizing population, showing that these packs were founded by multiple, unrelated wolves from Canada. High dispersal rates, long dispersal distances, and lack of a founding population bottleneck indicate that wolves in the United States and Canada should be viewed and managed as a single population. Restoration in the United States by artificial transplants from Alberta to Yellowstone National Park and central Idaho began in 1995. The transplanted wolves will likely aid demographic recovery, but permanently retaining the high genetic variation of wolves in the United States will require assuring gene flow throughout the central Rocky Mountains.     

Molecular Genetics of Pre-1940 Red Wolves

Conservation of the endangered red wolf (Canis rufus) has become a controversial issue because its genetic and morphological composition has been altered by hybridization with coyotes (C. latrans) and possibly gray wolves (C. lupus) making its evolutionary origins difficult to ascertain. The evolutionary hypothesis based on morphological data is that the red wolf had an Early Pleistocene origin and was the predecessor of both modern coyotes and gray wolves. After 1940 red wolves hybridized with coyotes as the species vanished from the wild. In contrast to this ancient origin-recent introgression hypothesis, molecular data are more consistent with an origin through hybridization between gray wolves and coyotes. Interspecific hybridization may have occurred repeatedly over time prior to European settlement in the southcentral United States or may have been induced recently by anthropogenic changes. We review recent molecular evidence and present new results from the analysis of mitochondrial and nuclear DNA markers in pre-1940 populations of red wolves. Our results are inconsistent with an ancient origin of the red wolf and support the hybridization model. We discuss possible hybridization scenarios and reasons for the red wolf reintroduction program to be concerned with the effects of genetic introgression from coyotes.     

A study of the genetic relationships within and among wolf packs using DNA fingerprinting and mitochondrial DNA

Summary DNA fingerprinting and mitochondrial DNA analyses have not been used in combination to study relatedness in natural populations. We present an approach that involves defining the mean fingerprint similarities among individuals thought to be unrelated because they have different mtDNA genotypes. Two classes of related individuals are identified by their distance in standard errors above this mean value. The number of standard errors is determined by analysis of the association between fingerprint similarity and relatedness in a population with a known genealogy. We apply this approach to gray wolf packs from Minnesota, Alaska, and the Northwest Territories. Our results show that: (1) wolf packs consist primarily of individuals that are closely related genetically, but some packs contain unrelated, non-reproducing individuals; (2) dispersal among packs within the same area is common; and (3) short-range dispersal appears more common for female than male wolves. The first two of these genetically-based observations are consistent with behavioral data on pack structure and dispersal in wolves, while the apparent sex bias in dispersal was not expected.USA Offprint requests to: N. Lehman at the present address     

Population structure of red drum (Sciaenops ocellatus) in the northern Gulf of Mexico, as inferred from variation in nuclear-encoded microsatellites

Allelic variation at eight nuclear-encoded microsatellites was assayed among 967 red drum (Sciaenops ocellatus) sampled from four consecutive cohorts at seven geographic localities (=28 samples total) in the northern Gulf of Mexico (Gulf). Number of alleles per microsatellite ranged from 6 to 21; average direct-count heterozygosity values per sample (-SE) ranged from 0.560ǂ.018 to 0.903ǂ.009. Tests of Hardy-Weinberg equilibrium revealed significant departures from expected genotype proportions at one microsatellite, which was omitted from further analysis. Tests of genotypic equilibrium indicated that genotypes between pairs of microsatellites were randomly associated. Homogeneity tests of allele distributions across cohorts within localities were non-significant following correction for multiple tests executed simultaneously, and results from molecular analysis of variance indicated that the genetic variance component attributable to variation among cohorts did not differ significantly from zero. Homogeneity tests of allele distributions among localities (cohorts pooled) revealed significant differences both before and after correction for multiple tests. Neighbor-joining clustering of a pairwise matrix of Š values (an unbiased estimator of F_ST), spatial autocorrelations, and regression analysis revealed a pattern of isolation by distance, where genetic divergence among geographic samples increases with geographic distance between sample localities. The pattern and degree of temporal and spatial divergence in the nuclear-encoded microsatellites paralleled almost exactly those of mitochondrial (mt) DNA, as determined in a prior study. Stability of both microsatellite and mtDNA allele distributions within localities indicates that the small but significant genetic divergence among geographic samples represents true signal and that overlapping populations of red drum in the northern Gulf may be influenced by independent population dynamics. The degree of genetic divergence in microsatellites and mtDNA is virtually identical, indicating that genetic effective size of microsatellites and mtDNA in red drum are the same. This, in turn, suggests either that gene flow in red drum in the northern Gulf could be biased sexually or that red drum populations may not be in equilibrium between genetic drift and migration. If a sexual bias exists, the observation that divergence in mtDNA is considerably less than 4 times that of microsatellites could suggest female-mediated dispersal and/or male philopatry. The observed isolation-by-distance effect indicates a practical limit to dispersal. Approximate estimates of geographic neighborhood size suggest the limit is in the range 700-900 km. Although the genetic studies of red drum indicate significant genetic divergence across the northern Gulf, the genetic differences do not delimit specific populations or stocks with fixed geographic boundaries.     

Demographic Responses by Birds to Forest Fragmentation

Abstract:  Despite intensive recent research on the effects of habitat loss and fragmentation on bird populations, our understanding of underlying demographic causes of population declines is limited. We reviewed avian demography in relation to habitat fragmentation. Then, through a meta-analysis, we compared specific demographic responses by forest birds to habitat fragmentation, providing a general perspective of factors that make some species and populations more vulnerable to fragmentation than others. We obtained data from the scientific literature on dispersal, survival, fecundity, and nesting success of birds. Birds were divided into subgroups on the basis of region, nest site, biogeographical history, and migration strategy. Species most sensitive to fragmentation were ground- or open-nesters nesting in shrubs or trees. Residents were equally sensitive to fragmentation in the Nearctic and Palearctic regions, but Nearctic migrants were more sensitive than Palearctic migrants. Old World species were less sensitive than New World species, which was predicted based on the history of forest fragmentation on these two continents. Pairing success was the variable most associated with fragmentation, suggesting an important role of dispersal. Fledgling number or condition, timing of nesting, and clutch size were not associated with sensitivity to fragmentation, suggesting that negative fragmentation effects on birds do not generally result from diminished food resources with increasing level of fragmentation. Future studies on demographic responses of birds to habitat fragmentation would be more effective if based on a combination of measures that can distinguish among the demographic mechanisms underlying population changes related to habitat fragmentation.     

Hybridization between Wolves and Dogs

Concern has been expressed that European populations of gray wolves ( Canis lupus ) have extensively hybridized with domestic dogs ( C. familiaris ). We reviewed and analyzed surveys of mitochondrial and biparentally inherited genetic markers in dogs and wild populations of wolf-like canids. Although dog-wolf hybrids have been observed in the wild, significant introgression of dog markers into wild wolf populations has not yet occurred. Our investigation suggests that hybridization may not be an important conservation concern even in small, endangered wolf populations near human settlements. The behavioral and physiological differences between domestic dogs and gray wolves may be sufficiently great such that mating is unlikely and hybrid offspring rarely survive to reproduce in the wild.     

Wolves-coyotes-foxes: a cascade among carnivores

Due to the widespread eradication of large canids and felids, top predators in many terrestrial ecosystems are now medium-sized carnivores such as coyotes. Coyotes have been shown to increase songbird and rodent abundance and diversity by suppressing populations of small carnivores such as domestic cats and foxes. The restoration of gray wolves to many parts of North America, however, could alter this interaction chain. Here we use a 30-year time series of wolf, coyote, and fox relative abundance from the state of Minnesota, USA, to show that wolves suppress coyote populations, which in turn releases foxes from top-down control by coyotes. In contrast to mesopredator release theory, which has often considered the consequence of top predator removal in a three-species interaction chain (e.g., coyote-fox-prey), the presence of the top predator releases the smaller predator in a four-species interaction chain. Thus, heavy predation by abundant small predators might be more similar to the historical ecosystem before top-predator extirpation. The restructuring of predator communities due to the loss or restoration of top predators is likely to alter the size spectrum of heavily consumed prey with important implications for biodiversity and human health.     

Indirect effects and traditional trophic cascades: a test involving wolves, coyotes, and pronghorn

The traditional trophic cascades model is based on consumer resource interactions at each link in a food chain. However, trophic-level interactions, such as mesocarnivore release resulting from intraguild predation, may also be important mediators of cascades. From September 2001 to August 2004, we used spatial and seasonal heterogeneity in wolf distribution and abundance in the southern Greater Yellowstone Ecosystem to evaluate whether mesopredator release of coyotes (Canis latrans), resulting from the extirpation of wolves (Canis lupus), accounts for high rates of coyote predation on pronghorn (Antilocapra americana) fawns observed in some areas. Results of this ecological perturbation in wolf densities, coyote densities, and pronghorn neonatal survival at wolf-free and wolf-abundant sites support the existence of a species-level trophic cascade. That wolves precipitated a trophic cascade was evidenced by fawn survival rates that were four-fold higher at sites used by wolves. A negative correlation between coyote and wolf densities supports the hypothesis that interspecific interactions between the two species facilitated the difference in fawn survival. Whereas densities of resident coyotes were similar between wolf-free and wolf-abundant sites, the abundance of transient coyotes was significantly lower in areas used by wolves. Thus, differential effects of wolves on solitary coyotes may be an important mechanism by which wolves limit coyote densities. Our results support the hypothesis that mesopredator release of coyotes contributes to high rates of coyote predation on pronghorn fawns, and demonstrate the importance of alternative food web pathways in structuring the dynamics of terrestrial systems.     

The importance of considering multiple interacting species for conservation of species at risk

Conservation of species at risk of extinction is complex and multifaceted. However, mitigation strategies are typically narrow in scope, an artifact of conservation research that is often limited to a single species or stressor. Knowledge of an entire community of strongly interacting species would greatly enhance the comprehensiveness and effectiveness of conservation decisions. We investigated how camera trapping and spatial count models, an extension of spatial-recapture models for unmarked populations, can accomplish this through a case study of threatened boreal woodland caribou (Rangifer tarandus caribou). Population declines in caribou are precipitous and well documented, but recovery strategies focus heavily on control of wolves (Canis lupus) and pay less attention to other known predators and apparent competitors. Obtaining necessary data on multispecies densities has been difficult. We used spatial count models to concurrently estimate densities of caribou, their predators (wolf, black bear [Ursus americanus], and coyote [Canis latrans]), and alternative prey (moose [Alces alces] and white-tailed deer [Odocoileus virginianus]) from a camera-trap array in a highly disturbed landscape within northern Alberta's Oil Sands Region. Median densities were 0.22 caribous (95% Bayesian credible interval [BCI] = 0.08–0.65), 0.77 wolves (95% BCI = 0.26–2.67), 2.39 moose (95% BCI = 0.56–7.00), 2.64 coyotes (95% BCI = 0.45–6.68), and 3.63 black bears (95% BCI = 1.25–8.52) per 100 km². (The white-tailed deer model did not converge.) Although wolf densities were higher than densities recommended for caribou conservation, we suggest the markedly higher black bear and coyote densities may be of greater concern, especially if government wolf control further releases these species. Caribou conservation with a singular focus on wolf control may leave caribou vulnerable to other predators. We recommend a broader focus on the interacting species within a community when conserving species.     

Genetic Structure and Migration in Native and Reintroduced Rocky Mountain Wolf Populations

Gray wolf (Canis lupus) recovery in the Rocky Mountains of the U.S. is proceeding by both natural recolonization and managed reintroduction. We used DNA microsatellite analysis of wolves transplanted from Canada to two reintroduction sites in the U.S. to study population structure in native and reintroduced wolf populations. Gene flow due to migration between regions in Canada is substantial, and all three recovery populations in the U.S. had high genetic variation. The reintroduced founders were moderately genetically divergent from the naturally colonizing U.S. population. These findings corroborate that the reintroduction more than meets generally accepted genetic guidelines. Maintaining this variation, however, will depend on ample reproduction in the first few generations. In the long term genetic variation will best be retained if migration occurs among the recolonizing and the two transplanted populations. Evidence from field observation and genetic studies shows extensive dispersal by wolves, and we conclude that exchange among these groups due to natural dispersal is likely if public tolerance and legal protection are adequate outside lands designated for wolf recovery.     

Longitudinal Analysis of Attitudes Toward Wolves

Understanding individual attitudes and how these predict overt opposition to predator conservation or direct, covert action against predators will help to recover and maintain them. Studies of attitudes toward wild animals rely primarily on samples of individuals at a single time point. We examined longitudinal change in individuals’ attitudes toward gray wolves (Canis lupus). In the contiguous United States, amidst persistent controversy and opposition, abundances of gray wolves are at their highest in 60 years. We used mailed surveys to sample 1892 residents of Wisconsin in 2001 or 2004 and then resampled 656 of these individuals who resided in wolf range in 2009. Our study spanned a period of policy shifts and increasing wolf abundance. Over time, the 656 respondents increased agreement with statements reflecting fear of wolves, the belief that wolves compete with hunters for deer (Odocoileus virginianus), and inclination to poach a wolf. Endorsement of lethal control of wolves by the state and public hunting of wolves also increased. Neither the time span over which respondents reported exposure to wolves locally nor self‐reported losses of domestic animals to wolves correlated with changes in attitude. We predict future increases in legal and illegal killing of wolves that may reduce their abundance in Wisconsin unless interventions are implemented to improve attitudes and behavior toward wolves. To assess whether interventions change attitudes, longitudinal studies like ours are needed. Análisis Longitudinal de las Actitudes Hacia Lobos     

Do yellow-bellied marmots respond to predator vocalizations?

We conducted four experiments to determine whether yellow-bellied marmots, Marmota flaviventris, discriminate among predator vocalizations, and if so, whether the recognition mechanism is learned or experience-independent. First, we broadcast to marmots the social sounds of coyotes, Canis latrans, wolves, Canis lupus, and golden eagles, Aquila chrysaetos, as well as conspecific alarm calls. Coyotes and eagles are extant predators at our study site, while wolves have been absent since the mid-1930s. In three follow-up experiments, we reversed the eagle call and presented marmots with forward and reverse calls to control for response to general properties of call structure rather than those specifically associated with eagles, we tested for novelty by comparing responses to familiar and unfamiliar birds, and we tested for the duration of predator sounds by comparing a wolf howl (that was much longer than the coyote in the first experiment) with a long coyote howl of equal duration to the original wolf. Marmots suppressed foraging and increased looking most after presentation of the conspecific alarm call and least after that of the coyote in the first experiment, with moderate responses to wolf and eagle calls. Marmots responded more to the forward eagle call than the reverse call, a finding consistent with a recognition template. Marmots did not differentiate vocalizations from the novel and familiar birds, suggesting that novelty itself did not explain our results. Furthermore, marmots did not differentiate between a wolf howl and a coyote howl of equal duration, suggesting that the duration of the vocalizations played a role in the marmots’ response. Our results show that marmots may respond to predators based solely on acoustic stimuli. The response to currently novel wolf calls suggests that they have an experience-independent ability to identify certain predators acoustically. Marmots’ response to predator vocalizations is not unexpected because 25 of 30 species in which acoustic predator discrimination has been studied have a demonstrated ability to respond selectively to cues from their predators.     

The Status and Conservation of the Wolf in Gujarat and Rajasthan, India

Abstract: Populations of wolves were estimated for the states of Gujarat and Rajasthan, India, based on interviews and surveys. The wolf range is continuous. The total number of wolves in Gujarat is between 190 and 270 and in Rajasthan is between 253 and 350. Recommendations for a wolf conservation strategy include: (1) encouraging public support and education, (2) enforcing legal protection, (3) paying compensation for wolf-killed livestock, (4) conducting surveys of wolf populations and research on the dynamics of select populations, (5) protecting breeding habitats, and (6) eradicating feral dogs from wolf conservation areas. It is not too late to save the wolf if such a conservation strategy is implemented immediately.     

Implications of Macroalgal Isolation by Distance for Networks of Marine Protected Areas

The global extent of macroalgal forests is declining, greatly affecting marine biodiversity at broad scales through the effects macroalgae have on ecosystem processes, habitat provision, and food web support. Networks of marine protected areas comprise one potential tool that may safeguard gene flow among macroalgal populations in the face of increasing population fragmentation caused by pollution, habitat modification, climate change, algal harvesting, trophic cascades, and other anthropogenic stressors. Optimal design of protected area networks requires knowledge of effective dispersal distances for a range of macroalgae. We conducted a global meta‐analysis based on data in the published literature to determine the generality of relation between genetic differentiation and geographic distance among macroalgal populations. We also examined whether spatial genetic variation differed significantly with respect to higher taxon, life history, and habitat characteristics. We found clear evidence of population isolation by distance across a multitude of macroalgal species. Genetic and geographic distance were positively correlated across 49 studies; a modal distance of 50–100 km maintained F_ST < 0.2. This relation was consistent for all algal divisions, life cycles, habitats, and molecular marker classes investigated. Incorporating knowledge of the spatial scales of gene flow into the design of marine protected area networks will help moderate anthropogenic increases in population isolation and inbreeding and contribute to the resilience of macroalgal forests. Implicaciones del Aislamiento por Distancia de Macroalgas para Redes de Áreas Marinas Protegidas     

Allozyme and mitochondrial DNA variation in Cuban populations of the shrimp Farfantepenaeus notialis (Crustacea: Decapoda)

We investigated the genetic diversity among populations of the shrimp Farfantepenaeus notialis, the most abundant penaeid species around Cuba. A total of 25 allozyme loci were analyzed in samples of shrimps from seven localities at the south central platform of the island (Ana María Gulf). Samples from three of these localities and from Batabanó Gulf and Guacanayabo Gulf at the south west and south east platforms of the island, respectively, were also characterized at the mtDNA level through sequence variation of a 2027 bp segment including part of the COI and COIII genes. Of the 25 allozyme loci studied 9 were polymorphic: Akp2, Akp3, AmyB, Est3, Gdh, GP7, and Per1, 2 and 3. In contrast to mtDNA, the pattern of allozyme variation among localities revealed strong population structuring at Ana María Gulf, with significant F _st in all pairwise comparisons. The magnitude of F _st estimates as well as the grouping pattern obtained by a UPGMA analysis based on a distance matrix indicated that the level of differentiation was concordant with the geographical position of the localities and the hydrographic regime. Homogeneity of mtDNA suggested that differentiation of allozyme loci might be due to more recent events rather than historical isolation of the sampled populations. Ana María and Guacanayabo Gulf populations were differentiated by mtDNA from Batabanó Gulf, at the southwestern end of the island. The analysis showed three restriction site differences among them, suggesting genetic isolation of the two regions. The present results also suggest that an artificial introduction of larvae from Tunas de Zaza into Batabanó Gulf, in an effort to repopulate this fishing region, may have been ineffective. Received: 13 December 1999 / Accepted: 2 October 2000     

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.     

Effects of Habitat Fragmentation on Effective Population Size in the Endangered Rio Grande Silvery Minnow

Abstract:  We assessed spatial and temporal patterns of genetic diversity to evaluate effects of river fragmentation on remnant populations of the federally endangered Rio Grande silvery minnow ( Hybognathus amarus ). Analysis of microsatellite and mitochondrial DNA detected little spatial genetic structure over the current geographic range, consistent with high gene flow despite fragmentation by dams. Maximum-likelihood analysis of temporal genetic data indicated, however, that present-day effective population size ( N_eV ) of the largest extant population of this species was 78 and the ratio of effective size to adult numbers ( N_eV/N ) was ∼ 0.001 during the study period (1999 to 2001). Coalescent-based analytical methods provided an estimate of historical (river fragmentation was completed in 1975) effective size ( N_eI  ) that ranged between 10⁵ and 10⁶. We propose that disparity between contemporary and historical estimates of N_e and low contemporary N_e/N result from recent changes in demography related to river fragmentation. Rio Grande silvery minnows produce pelagic eggs and larvae subject to downstream transport through diversion dams. This life-history feature results in heavy losses of yearly reproductive effort to emigration and mortality, and extremely large variance in reproductive success among individuals and spawning localities. Interaction of pelagic early life history and river fragmentation has altered demographic and genetic dynamics of remnant populations and reduced N_e to critically low values over ecological time.     

Dynamics of Hybridization and Introgression in Red Wolves and Coyotes

Abstract:  Hybridization and introgression are significant causes of endangerment in many taxa and are considered the greatest biological threats to the reintroduced population of red wolves ( Canis rufus ) in North Carolina (U.S.A.). Little is known, however, about these processes in red wolves and coyotes ( C. latrans ). We used individual-based simulations to examine the process of hybridization and introgression between these species. Under the range of circumstances we considered, red wolves in colonizing and established populations were quickly extirpated, persisted near the carrying capacity, or had intermediate outcomes. Sensitivity analyses suggested that the probabilities of quasi extinction and persistence of red wolves near the carrying capacity were most affected by the strength of two reproductive barriers: red wolf challenges and assortative mating between red wolves and coyotes. Because model parameters for these barriers may be difficult to estimate, we also sought to identify other predictors of red wolf population fate. The proportion of pure red wolves in the population was a strong predictor of the future probabilities of red wolf quasi extinction and persistence. Finally, we examined whether sterilization can be effective in minimizing introgression while allowing the reintroduced red wolf population to grow. Our results suggest sterilization can be an effective short-term strategy to reduce the likelihood of extirpation in colonizing populations of red wolves. Whether red wolf numbers are increased by sterilization depends on the level of sterilization effort and the acting reproductive barriers. Our results provide an outline of the conditions likely required for successful reestablishment and long-term maintenance of populations of wild red wolves in the presence of coyotes. Our modeling approach may prove generally useful in providing insight into situations involving complex species interactions when data are few.     

Biological, Conservation, and Ethical Implications of Exploiting and Controlling Wolves

The widespread claim that wolf populations can withstand 25–50% or greater annual reductions without major biological consequences is based primarily on the observation that populations often maintain their size from year to year as harvest or control continues or recover within a few years afterward. This emphasis on numerical status overlooks the likelihood of major, lingering impacts on the size, number, stability, and persistence of family-group social units, on reproductive, hunting, and territorial behavior, on the role of learning and related traditions, on within- and between-group patterns of genetic variation, and on overall mortality rates. The tendency of biologists and agencies in northern North America to promote wolf harvests that are four to eight times greater than ungulate harvests, in accord with the wolf versus ungulate difference in reproductive rates but contradictory to a broad array of differences in social organization and related behavior, is reason enough to question the logic of this prevailing management view. True sustained-yield management requires more emphasis on qualitative biological features to determine the extent to which wolves and other species with evolutionary histories as predators rather than as prey should be harvested. Most recent government-sponsored wolf control programs and proposals, including sterilization, relocation, and "redirected" killing, have been based on questionable claims about ungulate or livestock problems and have not adequately considered potential biological costs (especially to the target wolf populations), benefits, or management alternatives. The high sentience of wolves justifies overlapping biological-ethical concerns about such programs and especially about the heavy, indiscriminate, deceptively reported public hunting and trapping of wolves that is currently permitted throughout most of Alaska (U.S.A.)—including in national parks—and elsewhere.