Habitat preferences in gray marmots (Marmota baibacina)

We examined habitat preferences of the Southern Altai subspecies of gray marmots (Marmota baibacina baibacina) both at the small and large scale. Considerable differences in habitat use among the gray marmot (sub)species complex have been described; Marmota kastschenkoi possibly represents the only forest-dwelling Palearctic marmot. Our results show that habitat use in Southern Altai marmots is determined mainly by vegetation type. The Altai marmots preferred grasslands and shrublands and their distribution was limited to the alpine zone above timberline. Marmots clearly avoided woodlands, even the forest edges and forest-steppe areas with a tree cover greater than 10 %. Gray marmots occur rarely in habitats occupied by alpine pikas, whereas presence of ground squirrels had no effect on marmot distribution. Altai marmots preferred mesic habitats with permeable subsoil layers. Livestock grazing and human disturbance did not affect marmot occurrence. Habitat requirements of examined Altai subspecies M. baibacina baibacina differ from the forest-steppe M. kastschenkoi; nevertheless, the ecological factors to which the forest-dwelling species responds remain to be analyzed. A recent speciation process in gray marmot complex was followed by the evolution of ecological requirements resulting in adaptation to forest dwelling.     

Genetic and morphometric analysis of sixteenth century <Emphasis Type="Italic">Canis</Emphasis> skull fragments: implications for historic eastern and gray wolf distribution in North America

Resolving the taxonomy and historic ranges of species are essential to recovery plans for species at risk and conservation programs that aim to restore extirpated populations. In eastern North America, planning for wolf population restoration is complicated by the disputed historic distributions of two wolf species: the Old World-evolved gray wolf (Canis lupus) and the New World-evolved eastern wolf (C. lycaon). We used genetic and morphometric data from 4- to 500-year-old Canis samples excavated in London, Ontario, Canada to help clarify the historic range of these two wolf species in the eastern temperate forests of North America. We isolated DNA and sequenced the mitochondrial control region and found that none of the samples were of gray wolf origin. Two of the DNA sequences corresponded to those found in present day coyotes (C. latrans), but morphometric comparisons show an eastern wolf, not coyote, origin. The remaining two sequences matched ancient domestic dog haplotypes. These results suggest that the New World-evolved eastern wolf, not the gray wolf, occupied this region prior to the arrival of European settlers, although eastern-gray wolf hybrids cannot be ruled out. Furthermore, our data support the idea of a shared common ancestry between eastern wolves and western coyotes, and that the distribution of gray wolves at this time probably did not include the eastern temperate forests of North America.     

Quantifying Economic Impacts of Large-Carnivore Depredation on Bovine Calves

Abstract: We examined and quantified the economic impact of grizzly bear (Ursus arctos) and gray wolf (Canis lupus) depredation on calves in the Upper Green River Cattle Allotment in western Wyoming, USA, using records of the number of animals grazed and number lost during 1990–2004. Our analysis indicated that increased calf losses coincided with grizzly bear and gray wolf arrival and population establishment, with the first confirmed depredation by grizzly bears in 1995 and the first confirmed wolf depredation in 2000. From 1995 through 2004, 29,693 calves grazed on the allotment, and of the 1,332 calves lost to all causes, an estimated 520 calves were lost to grizzly bear depredation and 177 calves to gray wolf depredation. We examined past and current grizzly and gray wolf compensation programs with respect to reimbursement of producers for costs associated with large-carnivore depredation. Estimated 1995–2004 uncompensated financial impacts from grizzly bear and gray wolf calf losses on the allotment were US222,500. Our analysis suggested equitable compensation factors of 3.8:1 for grizzly bear depredation and 6.3:1 for gray wolf depredation. Inadequate compensation for livestock depredation results in resistance to large-carnivore recovery programs. Development of compensation programs that fairly reimburse livestock producers for losses is, therefore, a necessary component of carnivore recovery efforts. Our analysis also suggested that grizzly bear management actions were effectively targeting livestock-depredating grizzly bears on the allotment.     

Y‐chromosome evidence supports widespread signatures of three‐species Canis hybridization in eastern North America

There has been considerable discussion on the origin of the red wolf and eastern wolf and their evolution independent of the gray wolf. We analyzed mitochondrial DNA (mtDNA) and a Y‐chromosome intron sequence in combination with Y‐chromosome microsatellites from wolves and coyotes within the range of extensive wolf–coyote hybridization, that is, eastern North America. The detection of divergent Y‐chromosome haplotypes in the historic range of the eastern wolf is concordant with earlier mtDNA findings, and the absence of these haplotypes in western coyotes supports the existence of the North American evolved eastern wolf (Canis lycaon). Having haplotypes observed exclusively in eastern North America as a result of insufficient sampling in the historic range of the coyote or that these lineages subsequently went extinct in western geographies is unlikely given that eastern‐specific mtDNA and Y‐chromosome haplotypes represent lineages divergent from those observed in extant western coyotes. By combining Y‐chromosome and mtDNA distributional patterns, we identified hybrid genomes of eastern wolf, coyote, gray wolf, and potentially dog origin in Canis populations of central and eastern North America. The natural contemporary eastern Canis populations represent an important example of widespread introgression resulting in hybrid genomes across the original C. lycaon range that appears to be facilitated by the eastern wolf acting as a conduit for hybridization. Applying conventional taxonomic nomenclature and species‐based conservation initiatives, particularly in human‐modified landscapes, may be counterproductive to the effective management of these hybrids and fails to consider their evolutionary potential.     

A reappraisal of the evidence for regulation of wolf populations

The dogma that gray wolf (Canis lupus) population densities in naturally occurring systems are limited almost solely by available ungulate biomass is based upon studies that fit straight line linear regressions (Type 1 numerical response) to data collected at 32 sites across North America. We fit Type 1, 2, and 3 response functions to the data using linear and nonlinear regression as appropriate and found that the evidence supported wolf population regulation by density-dependence as much as limitation by prey availability. When we excluded 4 of 32 points from the original data set because those points represented exploited or expanding wolf populations the data suggested that wolf populations are self regulated rather than limited by prey biomass by at least a 3:1 margin. In establishing goals for sustainable wolf population levels, managers of wolf reintroductions and species recovery efforts should account for the possibility that some regulatory mechanism plays an important role in wolf population dynamics. © 2011 The Wildlife Society.     

Dispersal Behavior and the Connectivity Between Wolf Populations in Northern Europe

ABSTRACT The isolated gray wolf (Canis lupus) population of the Scandinavian Peninsular is suffering from inbreeding depression. We studied dispersal of 35 wolves fitted with very high frequency (20) or Global Positioning System—global system for mobile (15) radiocollars in the neighboring Finnish wolf population. The growing wolf population in Finland has high numbers of dispersing individuals that could potentially disperse into the Scandinavian population. About half (53%) of the dispersing wolves moved total distances that could have reached the Scandinavian population if they had been straight-line moves, but because of the irregular pattern of movements, we detected no wolves successfully dispersing to the Scandinavian population. Dispersal to the Scandinavian population was also limited by high mortality of wolves in reindeer (Rangifer tarandus) management areas and by dispersal to Bothnian Bay at times of the year when ice was not present. We suggest that when a small wolf population is separated from source populations by distance, barriers, and human exploitation, wildlife managers could promote the population's viability by limiting harvest in the peripheral areas or by introducing wolves from the source population.     

Population Viability, Nature Reserves, and the Outlook for Gray Wolf Conservation in North America

Theoretical work on population viability and extinction probabilities, empirical data from Canis lupus (gray wolf) populations, and expert opinion provide only general and conflicting conclusions about the number of wolves and the size of areas needed for conservation of wolf populations. There is no threshold population size or proven reserve design that guarantees long-term (century or more) survival for a gray wolf population. Most theoretical analyses of population viability have assumed a single, isolated population and lack of management intervention, neither of which is likely for wolves. Data on survival of actual wolf populations suggest greater resiliency than is indicated by theory. In our view, the previous theoretical treatments of population viability have not been appropriate to wolves, have contributed little to their conservation, and have created unnecessary dilemmas for wolf recovery programs by overstating the required population size. Nonetheless, viability as commonly understood may be problematic for small populations at the fringe of or outside the contiguous species range, unless they are part of a metapopulation. The capability of existing nature reserves to support viable wolf populations appears related to a variety of in situ circumstances, including size, shape and topography of the reserve; productivity, numbers, dispersion, and seasonal movement of prey; extent of poaching inside; degree of persecution outside; exposure to enzootica; attitudes of local people; and proximity to other wolf populations. We estimate that a population of 100 or more wolves and a reserve of several thousand square kilometers may be necessary to maintain a viable population in complete isolation, although 3000 km² or even 500–1000 km² may be adequate under favorable circumstances. In most cases, management intervention is probably necessary to assure the viability of relatively small, isolated populations. Because most reserves may be inadequate by themselves to ensure the long-term survival of wolf populations, favorable human attitudes toward the species and its management must be recognized as paramount, and cooperation of neighboring management jurisdictions will be increasingly important.     

Genetic nature of eastern wolves: Past, present and future

Eastern North American wolves have long been recognized as morphologically distinct from both coyotes and gray wolves. This has led to questions regarding their origins and taxonomic status. Eastern wolves are mainly viewed as: (1) a smaller subspecies of gray wolf (Canis lupus lycaon), potentially the result of historical hybridization between gray wolves (C. lupus) and red wolves (C. rufus), (2) a hybrid, the result of gray wolf (C. lupus) and coyote (C. latrans) interbreeding, or (3) a distinct species, C. lycaon, closely related to the red wolf (C. rufus). Although debate persists, recent molecular studies suggest that the eastern wolf is not a gray wolf subspecies, nor the result of gray wolf/coyote hybridization. Eastern wolves were more likely a distinct species, C. lycaon, prior to the eastward spread of coyotes in the late 1800s. However, contemporary interbreeding exits between C. lycaon to both C. lupus and C. latrans over much of its present range complicating its present taxonomic characterization. While hybridization may be reducing the taxonomic distinctiveness of C. lycaon, it should not necessarily be viewed as negative influence. Hybridization may be enhancing the adaptive potential of eastern wolves, allowing them to more effectively exploit available resources in rapidly changing environments.     

Efficient,Noninvasive Genetic Sampling for Monitoring Reintroduced Wolves

ABSTRACT Traditional methods of monitoring gray wolves (Canis lupus) are expensive and invasive and require extensive efforts to capture individual animals. Noninvasive genetic sampling (NGS) is an alternative method that can provide data to answer management questions and complement already-existing methods. In a 2-year study, we tested this approach for Idaho gray wolves in areas of known high and low wolf density. To focus sampling efforts across a large study area and increase our chances of detecting reproductive packs, we visited 964 areas with landscape characteristics similar to known wolf rendezvous sites. We collected scat or hair samples from 20% of sites and identified 122 wolves, using 8–9 microsatellite loci. We used the minimum count of wolves to accurately detect known differences in wolf density. Maximum likelihood and Bayesian single-session population estimators performed similarly and accurately estimated the population size, compared with a radiotelemetry population estimate, in both years, and an average of 1.7 captures per individual were necessary for achieving accurate population estimates. Subsampling scenarios revealed that both scat and hair samples were important for achieving accurate population estimates, but visiting 75% and 50% of the sites still gave reasonable estimates and reduced costs. Our research provides managers with an efficient and accurate method for monitoring high-density and low-density wolf populations in remote areas.     

Extinct Beringian wolf morphotype found in the continental U.S. has implications for wolf migration and evolution

Pleistocene diversity was much higher than today, for example there were three distinct wolf morphotypes (dire, gray, Beringian) in North America versus one today (gray). Previous fossil evidence suggested that these three groups overlapped ecologically, but split the landscape geographically. The Natural Trap Cave (NTC) fossil site in Wyoming, USA is an ideally placed late Pleistocene site to study the geographical movement of species from northern to middle North America before, during, and after the last glacial maximum. Until now, it has been unclear what type of wolf was present at NTC. We analyzed morphometrics of three wolf groups (dire, extant North American gray, Alaskan Beringian) to determine which wolves were present at NTC and what this indicates about wolf diversity and migration in Pleistocene North America. Results show NTC wolves group with Alaskan Beringian wolves. This provides the first morphological evidence for Beringian wolves in mid‐continental North America. Their location at NTC and their radiocarbon ages suggest that they followed a temporary channel through the glaciers. Results suggest high levels of competition and diversity in Pleistocene North American wolves. The presence of mid‐continental Beringian morphotypes adds important data for untangling the history of immigration and evolution of Canis in North America.     

Population structure and gene flow in a newly harvested gray wolf (<Emphasis Type="Italic">Canis lupus</Emphasis>) population

The genetic effects of harvest may be especially important in species that form social groups, such as gray wolves (Canis lupus). Though much research exists on the ecology and population dynamics of gray wolves, little research has focused on how anthropogenic harvest relates to the genetics of wolf populations. To analyze the short-term genetic consequences of the first two years of public wolf harvest in Minnesota following delisting under the Endangered Species Act, we genotyped harvested individuals at 18 microsatellite loci and quantified changes in population genetic structure and diversity in the first post-harvest year. If the harvest rate was high enough to create detectable genetic changes, population structure and differentiation between clusters could both increase because of decreased natal dispersal and increased disperser mortality, or they could decrease because of increased immigration from outside the population. In the Minnesota population, heterozygosity and allelic richness were not significantly different between years. However, population genetic structure increased and effective migration decreased among the sampled wolves. While the role of anthropogenic harvest in these changes cannot be distinguished from other confounding factors, this analysis suggests that harvest has a non-negligible effect and indicates the need for continued study to determine whether harvest-induced changes in genetic structure affect the evolutionary trajectory of harvested populations.     

Surveying Predicted Rendezvous Sites to Monitor Gray Wolf Populations

ABSTRACT We used rendezvous site locations of wolf (Canis lupus) packs recorded during 1996–2006 to build a predictive model of gray wolf rendezvous site habitat in Idaho, USA. Variables in our best model included green leaf biomass (Normalized Difference Vegetation Index), surface roughness, and profile curvature, indicating that wolves consistently used wet meadow complexes for rendezvous sites. We then used this predictive model to stratify habitat and guide survey efforts designed to document wolf pack distribution and fecundity in 4 study areas in Idaho. We detected all 15 wolf packs (32 wolf pack-yr) and 20 out of 27 (74%) litters of pups by surveying <11% of the total study area. In addition, we were able to obtain detailed observations on wolf packs (e.g., hair and scat samples) once we located their rendezvous sites. Given an expected decrease in the ability of managers to maintain radiocollar contact with all of the wolf packs in the northern Rocky Mountains, rendezvous sites predicted by our model can be the starting point and foundation for targeted sampling and future wolf population monitoring surveys.     

Large‐scale patterns of seed removal by small mammals differ between areas of low‐ versus high‐wolf occupancy

Because most tree species recruit from seeds, seed predation by small‐mammal granivores may be important for determining plant distribution and regeneration in forests. Despite the importance of seed predation, large‐scale patterns of small‐mammal granivory are often highly variable and thus difficult to predict. We hypothesize distributions of apex predators can create large‐scale variation in the distribution and abundance of mesopredators that consume small mammals, creating predictable areas of high and low granivory. For example, because gray wolf (Canis lupus) territories are characterized by relatively less use by coyotes (C. latrans) and greater use by foxes (Vulpes vulpes, Urocyon cinereoargentus) that consume a greater proportion of small mammals, wolf territories may be areas of reduced small‐mammal granivory. Using large‐scale, multiyear field trials at 22 sites with high‐ and low‐wolf occupancy in northern Wisconsin, we evaluated whether removal of seeds of four tree species was lower in wolf territories. Consistent with the hypothesized consequences of wolf occupancy, seed removal of three species was more than 25% lower in high‐wolf‐occupancy areas across 2 years and small‐mammal abundance was more than 40% lower in high‐wolf areas during one of two study years. These significant results, in conjunction with evidence of seed consumption in situ and the absence of significant habitat differences between high‐ and low‐wolf areas, suggest that top‐down effects of wolves on small‐mammal granivory and seed survival may occur. Understanding how interactions among carnivores create spatial patterns in interactions among lower trophic levels may allow for more accurate predictions of large‐scale patterns in seed survival and forest composition.     

Canid hybridization: contemporary evolution in human‐modified landscapes

Contemporary evolution through human‐induced hybridization occurs throughout the taxonomic range. Formerly allopatric species appear especially susceptible to hybridization. Consequently, hybridization is expected to be more common in regions with recent sympatry owing to human activity than in areas of historical range overlap. Coyotes ( Canis latrans) and gray wolves ( C. lupus) are historically sympatric in western North America. Following European settlement gray wolf range contracted, whereas coyote range expanded to include eastern North America. Furthermore, wolves with New World (NW) mitochondrial DNA (mtDNA) haplotypes now extend from Manitoba to Québec in Canada and hybridize with gray wolves and coyotes. Using mtDNA and 12 microsatellite markers, we evaluated levels of wolf‐coyote hybridization in regions where coyotes were present (the Canadian Prairies, n = 109 samples) and absent historically (Québec, n = 154). Wolves with NW mtDNA extended from central Saskatchewan (51°N, 69°W) to northeastern Québec (54°N, 108°W). On the Prairies, 6.3% of coyotes and 9.2% of wolves had genetic profiles suggesting wolf‐coyote hybridization. In contrast, 12.6% of coyotes and 37.4% of wolves in Québec had profiles indicating hybrid origin. Wolves with NW and Old World ( C. lupus) mtDNA appear to form integrated populations in both regions. Our results suggest that hybridization is more frequent in historically allopatric populations. Range shifts, now expected across taxa following climate change and other human influence on the environment, might therefore promote contemporary evolution by hybridization.     

A case of visceral leishmaniosis in a gray wolf (Canis lupus) from Croatia

The southern habitats of Croatia's gray wolf (Canis lupus) population are found in central and southern parts of Dalmatia. This region is recognized as an endemic region for canine visceral leishmaniosis, caused by Leishmania infantum. In November 2003, a 4-yr-old male gray wolf was found dead in the northwestern border of this endemic region. Pathologic and parasitologic analysis, confirmed by polymerase chain reaction, indicated that lesions associated with infection by Leishmania infantum are, in this case, typical for visceral leshmaniosis commonly described in dogs. Review of the literature suggests that this is the first reported case of gray wolf death due to lesions caused by L. infantum.     

Wolves in the Great Lakes region: a phylogeographic puzzle

Empirical studies demonstrate that natural hybridization in animals is more common than thought so far ( Mallet 2005 ), particularly among species that originated recently through cycles of population contraction–expansion arising from climate changes over the last glacial period, the Pleistocene. In addition, the post‐glacial global growth of human populations has fostered anthropogenic hybridization events, mediated by habitat changes, the persecution of large predators and the introduction of alien species ( Allendorf et al. 2001 ). The Canis lineage shows cases of both natural and anthropogenic hybridization, exacerbating the controversy about the number of species that should be formally validated in the taxonomic lists, the evolutionary role of genetic introgression and the ways to manage hybrids with invading wild or domesticated populations. The study by Wheeldon et al. (2010) , published in this issue of Molecular Ecology, adds a new piece to the intricate puzzle of evolution and taxonomy of Canis in North America. They show that sympatric wolves (C. lupus) and coyotes (C. latrans) are not (extensively) hybridizing in the western North American Great Lakes region (GLR). Widespread hybridization between coyotes and a genetically distinct, but closely related, wolf‐like population (the eastern wolf) occurred in the northeastern regions of North America. In Wheeldon et al.’s (2010) opinion, these data should prove definitely that two different species of wolf (the western gray wolf C. lupus and the eastern wolf C. lycaon) and their hybrids are distributed across the GLR.     

Population-genetical analysis of Altai osmans (<Emphasis Type="Italic">Oreoleuciscus</Emphasis>, Cyprinidae) from water bodies of Mongolia

Similarity of populations of Altai osmans Oreoleuciscus (Cyprinidae) is estimated on the material from all principal parts of their area in Mongolia using methods of population-genetical analysis. Nineteen loci are considered coding isozymes and isoforms of non-enzyme proteins on the material of ten samples, RAPD-analysis is made for two samples representing different population sets. The analyzed populations of Altai osmans on the Mongolia territory are subdivided into two spatial and genetically clearly differentiated population groups—the Selenga group (with the Tesiyn-Gol basin) and the Central Asian group (populations from water bodies of Lake Valley and Great Lakes Basin. Probable causes and mechanisms of formation of the revealed population-genetical structure are discussed.     

Human dimensions of wolf (<Emphasis Type="Italic">Canis lupus</Emphasis>) conflicts in Finland

The growth of the Finnish wolf (Canis lupus) population during the last years has highlighted people’s contradictory attitudes toward wolves. The supranational conservation objectives brought on by Finland’s membership in the European Union (EU) and the regional application of the official policy on wolves have led to conflicts. This article is based on the preparation process of the wolf management plan in Finland. As a part of the process, a nationwide hearing was arranged in 2004. Fear of wolves is widespread and even more common in areas with low wolf population. People living in areas where wolves occur feel that they can no longer influence decision making which affects them and that the authorities, conservationists, and the EU do not listen to their opinions. Numerous requests have been made for the legislation and its interpretation, and concessions to these requests would promote consensus and increase tolerance for wolves. Proposed changes include reforming the damage compensation system and formulating a clearer interpretation of the conservation status of wolves. However, it seems impossible to create a policy that would be universally supported. As the wolf question has become more central in environmental policy, the conservation and management of the species have become more complicated, and the cooperation between various interest groups has declined. The wolf conflict could be mitigated by compromises, but because the conflict is value-based, solutions are illusive.     

Molecular genetics of the most endangered canid: the Ethiopian wolf Canis simensis

The world's most endangered canid is the Ethiopian wolf Canis simensis , which is found in six isolated areas of the Ethiopian highlands with a total population of no more than 500 individuals. Ethiopian wolf populations are declining due to habitat loss and extermination by humans. Moreover, in at least one population, Ethiopian wolves are sympatric with domestic dogs, which may hybridize with them, compete for food, and act as disease vectors. Using molecular techniques, we address four questions concerning Ethiopian wolves that have conservation implications. First, we determine the relationships of Ethiopian wolves to other wolf-like canids by phylogenetic analysis of 2001 base pairs of mitochondrial DNA (mtDNA) sequence. Our results suggest that the Ethiopian wolf is a distinct species more closely related to gray wolves and coyotes than to any African canid. The mtDNA sequence similarity with gray wolves implies that the Ethiopian wolf may hybridize with domestic dogs, a recent derivative of the gray wolf. We examine this possibility through mtDNA restriction fragment analysis and analysis of nine microsatellite loci in populations of Ethiopian wolves. The results imply that hybridization has occurred between female Ethiopian wolves and male domestic dogs in one population. Finally, we assess levels of variability within and between two Ethiopian wolf populations. Although these closely situated populations are not differentiated, the level of variability in both is low, suggesting long-term effective population sizes of less than a few hundred individuals. We recommend immediate captive breeding of Ethiopian wolves to protect their gene pool from dilution and further loss of genetic variability.     

Considerations for Developing Wolf Harvesting Regulations in the Contiguous United States

Abstract: As gray wolves (Canis lupus) are removed from the federal Endangered Species List, management reverts to the states. Eventually most states will probably allow public wolf harvesting. Open seasons between about 1 November and 1 March accord more with basic wolf biology than during other times. Managers who consider wolf biology and public sensitivities, adapt public-taking regulations accordingly, and adjust harvest regulations as they learn will be best able to maximize the recreational value of wolf harvesting, minimize public animosity toward it, and meet their harvest objectives.