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.     

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     

Mitochondrial DNA Variability of the Gray Wolf: Genetic Consequences of Population Decline and Habitat Fragmentation

The gray wolf is a large, highly mobile predator whose original geographic range included most of the Northern Hemisphere. High rates of genetic exchange probably characterized even distantly-separated populations in the past, but recent population declines and habitat fragmentation have isolated previously contiguous populations, especially in the Old World. We examine mitochondrial DNA (mtDNA) variability among twenty-six populations of wolves from throughout their geographic range. We find eighteen mtDNA genotypes in gray wolves, seven of these are derived from hybridization with coyotes, four are confined to the New World, six are confined to the Old World and one is shared by both areas. Genetic differentiation among wolf populations is significant but small in magnitude. In the Old World, most localities have a single unique genotype, whereas in the New World several genotypes occur at most localities and three of the five genotypes are nearly ubiquitous. The pattern of genetic differentiation in the gray wolf contrasts with that of another large, highly vagile canid, the coyote, in which genetic differentiation among populations is not significant even among widely separated localities. We suggest that the difference between these two species reflects the rapid, recent increase in coyote numbers and expansion of their geographic range, and the coincident decline in gray wolf populations. Apparent genetic differences among extant wolf populations may be a recent phenomenon reflecting population declines and habitat fragmentation rather than a long history of genetic isolation.     

Paying for Tolerance: Rural Citizens' Attitudes toward Wolf Depredation and Compensation

Abstract:   As wolf ( Canis lupus ) populations recover in Wisconsin (U.S.A.), their depredations on livestock, pets, and hunting dogs have increased. We used a mail-back survey to assess the tolerance of 535 rural citizens of wolves and their preferences regarding the management of "problem" wolves. Specifically, we tested whether people who had lost domestic animals to wolves or other predators were less tolerant of wolves than neighboring residents who had not and whether compensation payments improved tolerance of wolves. We assessed tolerance via proxy measures related to an individual's preferred wolf population size for Wisconsin and the likelihood she or he would shoot a wolf. We also measured individuals' approval of lethal control and other wolf-management tactics under five conflict scenarios. Multivariate analysis revealed that the strongest predictor of tolerance was social group. Bear ( Ursus americanus ) hunters were concerned about losing valuable hounds to wolves and were more likely to approve of lethal control and reducing the wolf population than were livestock producers, who were more concerned than general residents. To a lesser degree, education level, experience of loss, and gender were also significant. Livestock producers and bear hunters who had been compensated for their losses to wolves were not more tolerant than their counterparts who alleged a loss but received no compensation. Yet all respondents approved of compensation payments as a management strategy. Our results indicate that deep-rooted social identity and occupation are more powerful predictors of tolerance of wolves than individual encounters with these large carnivores.     

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.     

Desert pastoralists’ negative and positive effects on rare wildlife in the Gobi

In arid regions of the developing world, pastoralists and livestock commonly inhabit protected areas, resulting in human–wildlife conflict. Conflict is inextricably linked to the ecological processes shaping relationships between pastoralists and native herbivores and carnivores. To elucidate relationships underpinning human–wildlife conflict, we synthesized 15 years of ecological and ethnographic data from Ikh Nart Nature Reserve in Mongolia's Gobi steppe. The density of argali (Ovis ammon), the world's largest wild sheep, at Ikh Nart was among the highest in Mongolia, yet livestock were >90% of ungulate biomass and dogs >90% of large‐carnivore biomass. For argali, pastoral activities decreased food availability, increased mortality from dog predation, and potentially increased disease risk. Isotope analyses indicated that livestock accounted for >50% of the diet of the majority of gray wolves (Canis lupus) and up to 90% of diet in 25% of sampled wolves (n = 8). Livestock composed at least 96% of ungulate prey in the single wolf pack for which we collected species‐specific prey data. Interviews with pastoralists indicated that wolves annually killed 1–4% of Ikh Nart's livestock, and pastoralists killed wolves in retribution. Pastoralists reduced wolf survival by killing them, but their livestock were an abundant food source for wolves. Consequently, wolf density appeared to be largely decoupled from argali density, and pastoralists had indirect effects on argali that could be negative if pastoralists increased wolf density (apparent competition) or positive if pastoralists decreased wolf predation (apparent facilitation). Ikh Nart's argali population was stable despite these threats, but livestock are increasingly dominant numerically and functionally relative to argali. To support both native wildlife and pastoral livelihoods, we suggest training dogs to not kill argali, community insurance against livestock losses to wolves, reintroducing key native prey species to hotspots of human–wolf conflict, and developing incentives for pastoralists to reduce livestock density.     

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.     

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.     

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.     

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.     

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     

The Challenge and Opportunity of Recovering Wolf Populations

The gray wolf once inhabited a wide variety of habitats throughout most of the northern hemisphere north of 20°N latitude. Because the animal preyed on livestock and competed with humans for wild prey, it was extirpated from much of its range outside of wilderness areas. Environmental awareness in the late 1960s brought for the wolf legal protection, increased research, and favorable media coverage. The species has increased in both Europe and North America, is beginning to reoccupy semiwilderness and agricultural land, and is causing increased damage to livestock. Because of the wolf's high reproductive rate and long dispersal tendencies, the animal can recolonize many more areas. In most such areas control will be necessary, but the same public sentiments that promoted wolf recovery reject control. If wolf advocates could accept control by the public rather than by the government, wolves could live in far more places. Insistence on government control discourages some officials and government agencies from promoting recovery. The use of large- or small-scale zoning for wolf management may help resolve the issue. Public education is probably the most effective way to minimize the problem and maximize wolf recovery, but the effort must begin immediately.     

Tracking a half century of media reporting on gray wolves

Natural resource and wildlife managers must balance the disparate priorities of a diversity of stakeholders. To manage these priorities, a firm understanding of topics salient to the public is needed. The media often report on issues of importance to the public; therefore, these reports may be a useful measure of public interest. However, efficient methods for distinguishing diverse topics related to a wildlife management issue reported in the media and changes in the salience of those topics have been lacking. We used latent Dirichlet allocation, a Bayesian mixture model, to quantitatively assess the salience of topics surrounding the gray wolf (Canis lupus), which was reintroduced to Idaho (U.S.A.) in 1995. We analyzed articles published from 1960 to 2015 in an Idaho newspaper. We identified 6 distinct topics associated with gray wolves: policy, hunting, biological status, implementation of management, recovery, and human-wolf conflict. The salience of topics pre- and postreintroduction of wolves (1995) and pre- and postdelisting of wolves from the U.S. Endangered Species Act (2009) differed significantly, underscoring that these events were turning points in how issues were being publicly discussed and framed. Articles written by the local reporters were more likely to report on topics regarding conflict between humans and wolves, whereas articles sourced from a national outlet reported more on topics pertaining to wolf policy and biological status. In the context of managing a contentious, far-ranging, and long-lived wildlife species, our methods can help guide the location and timing of a suite of management strategies (e.g., media relation plans and stakeholder engagement) that promote human-wildlife coexistence across different landscapes.     

Effects of Social Structure and Prey Dynamics on Extinction Risk in Gray Wolves

Extinction models based on diffusion theory generally fail to incorporate two important aspects of population biology—social structure and prey dynamics. We include these aspects in an individual-based extinction model for small, isolated populations of the gray wolf (Canis lupus). Our model predicts mean times to extinction significantly longer than those predicted by more general (diffusion) models. According to our model, an isolated population of 50 wolves has a 95% chance of surviving just 9 years and only a 30% chance of surviving beyond 100 years. Reflecting the influence of social structure, a wolf population initially comprising 50 individuals is expected to persist only a few years longer, on average (71 years), than is a population initially comprising just a single reproductive pair (62 years). In contrast, substantially greater average prey abundance leads to dramatically longer expected persistence times. Autocorrelated prey dynamics result in a more complex distribution of extinction times than predicted by many extinction models. We contend that demographic stochasticity may pose the greatest threat to small, isolated wolf populations, although environmental stochasticity and genetic effects may compound this threat. Our work highlights the importance of considering social structure and resource dynamics in the development of population viability analyses.     

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.     

Inbreeding Depression in a Captive Wolf (Canis lupus) Population

Abstract. Uncertainty currently exists regarding the extent to which mammalian carnivores suffer from inbreeding depression. In particular, it has been proposed that wolves and species with a similar social structure are adapted to close inbreeding. Empirical data, however, are scarce. This paper provides strong evidence against the contention that natural populations of wolves are resistant to inbreeding depression. We analyzed studbook data of a captive wolf population bred in Scandinavian zoos and found negative effects of inbreeding expressed as reductions in juvenile weight, reproduction, and longevity. The occurrence of an apparently bereditary form of blindness is also associated with inbreeding. Different effects of inbreeding can be attributed to genes originating from different founder pairs, thus indicating that alleles that are deleterious in the homozygous state are fairly common in natural wolf populations.     

Moral dimensions of human–wildlife conflict

Despite increasing support for conservation globally, controversy over specific conservation policies persists among diverse stakeholders. Investigating the links between morals in relation to conservation can help increase understanding about why humans support or oppose policy, especially related to human–wildlife conflict or human conflict over wildlife. Yet the moral dimension of human–wildlife conflict has mostly gone unconsidered and unmeasured; thus, policy and programmatic efforts to reduce controversy may be missing a key part of the equation. We conducted a web‐based survey (n = 1239 respondents) in Michigan (U.S.A.) to investigate cognitive and emotional influences on the value–behavior relationship. Respondents were identified by their interest and involvement in Michigan wolf management. The survey consisted of questions about values, emotions, cognitions, and behaviors relative to wolves in Michigan. We used path analysis to explore whether emotions and cognitions mediated the relationship between value and behavior. Most respondents attributed intrinsic value to wolves (n = 734) and all life (n = 773) and engaged in behaviors that benefited wolf populations and ecosystems regardless of stakeholder group (e.g., environmentalist, farmer). Attributing intrinsic value to wolves was positively related to favorable emotions toward wolves and cognitive assessments that hunting and trapping of wolves is unacceptable. Despite similarities in attribution of intrinsic value, groups differed in emotions and cognitions about wolf hunting. These differences provide a useful way to predict stakeholder behavior. Our findings may inform interventions aimed at increasing support for wolf management policies and positive interactions among stakeholders and wildlife. Leveraging agreement over intrinsic value may foster cooperation among stakeholders and garner support for controversial conservation policy.     

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.     

The causes and consequences of partial prey consumption by wolves preying on moose

For a wide range of taxa, partial prey consumption (PPC) is a frequent occurrence. PPC may arise from physiological constraints to gut capacity or digestive rate. Alternatively, PPC may represent an optimal foraging strategy. Assessments that clearly distinguish between these causes are rare and have been conducted only for invertebrate species that are ambush predators with extra-intestinal digestion (e.g., wolf spiders). We present the first strong test for the cause of PPC in a cursorial vertebrate predator with intestinal digestion: wolves (Canis lupus) feeding on moose (Alces alces). Previous theoretical assessments indicate that if PPC represents an optimal foraging strategy and is not caused by physiological limitations, then mean carcass utilization is negatively correlated with mean kill rate and the utilization of individual carcasses is uncorrelated with time between kills. Wolves exhibit exactly this pattern. We explore how the typical portrayal of PPC by wolves has been not only misleading but also detrimental to conservation by promoting negative attitudes toward wolves.     

Are wolves saving Yellowstone's aspen? A landscape-level test of a behaviorally mediated trophic cascade

Behaviorally mediated trophic cascades (BMTCs) occur when the fear of predation among herbivores enhances plant productivity. Based primarily on systems involving small-bodied predators, BMTCs have been proposed as both strong and ubiquitous in natural ecosystems. Recently, however, synthetic work has suggested that the existence of BMTCs may be mediated by predator hunting mode, whereby passive (sit-and-wait) predators have much stronger effects than active (coursing) predators. One BMTC that has been proposed for a wide-ranging active predator system involves the reintroduction of wolves (Canis lupus) to Yellowstone National Park, USA, which is thought to be leading to a recovery of trembling aspen (Populus tremuloides) by causing elk (Cervus elaphus) to avoid foraging in risky areas. Although this BMTC has been generally accepted and highly popularized, it has never been adequately tested. We assessed whether wolves influence aspen by obtaining detailed demographic data on aspen Stands using tree rings and by monitoring browsing levels in experimental elk exclosures arrayed across a gradient of predation risk for three years. Our study demonstrates that the historical failure of aspen to regenerate varied widely among stands (last recruitment year ranged from 1892 to 1956), and our data do not indicate an abrupt cessation of recruitment. This pattern of recruitment failure appears more consistent with a gradual increase in elk numbers rather than a rapid behavioral shift in elk foraging following wolf extirpation. In addition, our estimates of relative survivorship of young browsable aspen indicate that aspen are not currently recovering in Yellowstone, even in the presence of a large wolf population. Finally, in an experimental test of the BMTC hypothesis we found that the impacts of elk browsing on aspen demography are not diminished in sites where elk are at higher risk of predation by wolves. These findings suggest the need to further evaluate how trophic cascades are mediated by predator-prey life history and ecological context.