Invading white-tailed deer change wolf–caribou dynamics in northeastern Alberta

Human-caused habitat change has been implicated in current woodland caribou (Rangifer tarandus caribou) population declines across North America. Increased early seral habitat associated with industrial footprint can result in an increase in ungulate densities and subsequently those of their predator, wolves (Canis lupus). Higher wolf densities can result in increased encounters between wolves and caribou and consequently higher caribou mortality. We contrasted changes in moose (Alces alces) and deer (Odocoileus spp.) densities and assessed their effects on wolf–caribou dynamics in northeastern Alberta, Canada, pre (1994–1997) versus post (2005–2009) major industrial expansion in the region. Observable white-tailed deer (O. virginianus) increased 17.5-fold but moose remained unchanged. Wolf numbers also increased from approximately 6–11.5/1,000 km². Coincident with these changes, spatial overlap between wolf pack territories and caribou range was high relative to the mid-1990s. The high number of wolf locations in caribou range suggests that forays were not merely exploratory, but rather represented hunting forays and denning locations. Scat analysis indicated that wolf consumption of moose declined substantively during this time period, whereas use of deer increased markedly and deer replaced moose as the primary prey of wolves. Caribou increased 10-fold in the diet of wolves and caribou population trends in the region changed from stable to declining. Wolf use of beaver (Castor canadensis) increased since the mid-1990s. We suggest that recent declines in woodland caribou populations in the southerly extent of their range have occurred because high deer densities resulted in a numeric response by wolves and consequently higher incidental predation on caribou. Our results indicate that management actions to conserve caribou must now include deer in primary prey and wolf reduction programs. © 2010 The Wildlife Society     

Disturbance-Mediated Apparent Competition Decouples in a Northern Boreal Caribou Range

The most widely reported threat to boreal and mountain populations of woodland caribou (Rangifer tarandus caribou; caribou) involves habitat- or disturbance-mediated apparent competition (DMAC). With DMAC, natural and anthropogenic disturbances that increase the abundance of deciduous-browsing cervids (e.g., moose [Alces alces], deer [Odocoileus spp.]) are thought to promote predator (especially wolf [Canis lupus]) numbers, which heightens predation risk to caribou. We know most about the effects of DMAC on caribou where the species is under threat by anthropogenic activities in relatively productive southern boreal and mountain systems. Yet, >60% of extant boreal caribou range in North America consists of northern shield and taiga ecoregions of low productivity where caribou may compete with only 1 ungulate species (moose) in the context of DMAC. In this environment, we know very little of how DMAC acts as a limiting factor to caribou. In Saskatchewan, Canada, from 2014–2018, using a combination of vegetation sampling, aerial surveys, and telemetry data (n = 38 wolves), we searched for evidence of DMAC (trends in data consistent with the hypothesis) in an 87,193-km² section of the Western Boreal Shield, a poorly productive but pristine region (0.18% of land cover classed as an anthropogenic feature) with a historically high fire-return interval (47% of stands aged <40 years). Despite the high levels of disturbance, moose density was relatively low (47 moose/1,000 km²), likely because of the scarcity of deciduous or mixed-wood stands and low abundance of deciduous browse in the young conifer stands that dominated the landscape. In contrast, boreal caribou density was relatively high for the species (37 caribou/1,000 km²). Wolf density (3.1 wolves/1,000 km²) and pack sizes ( = 4.0 wolves/pack) were low and resident (established) territories were large ( = 4,360 km²; 100% minimum convex polygon). The low density of wolves mirrored the low (standardized) ungulate biomass index (UBI; moose + boreal caribou) of the study area (0.36 UBI/km²). We conclude that wolf and hence caribou populations were not responding in accordance with the outcomes generally predicted by DMAC in our study area because the requisite strong, positive response to fire of deciduous-browse and alternate-prey abundance was lacking. As a limiting factor to caribou, DMAC is likely modulated at a macroecological scale by factors such as net primary productivity, a corollary to the general hypothesis that we advance here (i.e., primary productivity hypothesis of DMAC). We caution against managing for caribou based on the presumption of DMAC where the mechanism does not apply, which may include much of boreal caribou range in the north. © 2020 The Wildlife Society.     

Wolves,white‐tailed deer,and beaver: implications of seasonal prey switching for woodland caribou declines

Population increases of primary prey can negatively impact alternate prey populations via demographic and behavioural responses of a shared predator through apparent competition. Seasonal variation in prey selection patterns by predators also can affect secondary and incidental prey by reducing spatial separation. Global warming and landscape changes in Alberta's bitumen sands have resulted in prey enrichment, which is changing the large mammal predator–prey system and causing declines in woodland caribou Rangifer tarandus caribou populations. We assessed seasonal patterns of prey use and spatial selection by wolves Canis lupus in two woodland caribou ranges in northeastern Alberta, Canada, that have undergone prey enrichment following recent white‐tailed deer Odocoileus virginianus invasion. We determined whether risk of predation for caribou (incidental prey) and the proportion of wolf‐caused‐caribou mortalities varied with season. We found that wolves showed seasonal variation in primary prey use, with deer and beaver Castor canadensis being the most common prey items in wolf diet in winter and summer, respectively. These seasonal dietary patterns were reflected in seasonal wolf spatial resource selection and resulted in contrasting spatial relationships between wolves and caribou. During winter, wolf selection for areas used by deer maintained strong spatial separation between wolves and caribou, whereas wolf selection for areas used by beaver in summer increased the overlap with caribou. Changing patterns in wolf resource selection were reflected by caribou mortality patterns, with 76.2% of 42 adult female caribou mortalities occurring in summer. Understanding seasonal patterns of predation following prey enrichment in a multiprey system is essential when assessing the effect of predation on an incidental prey species. Our results support the conclusion that wolves are proximately responsible for woodland caribou population declines throughout much of their range.     

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.     

Prey specialization may influence patterns of gene flow in wolves of the Canadian Northwest

This study characterizes population genetic structure among grey wolves (Canis lupus) in northwestern Canada, and discusses potential physical and biological determinants of this structure. Four hundred and ninety-one grey wolves, from nine regions in the Yukon, Northwest Territories and British Columbia, were genotyped using nine microsatellite loci. Results indicate that wolf gene flow is reduced significantly across the Mackenzie River, most likely due to the north-south migration patterns of the barren-ground caribou herds that flank it. Furthermore, although Banks and Victoria Island wolves are genetically similar, they are distinct from mainland wolf populations across the Amundsen Gulf. However, low-level island-mainland wolf migration may occur in conjunction with the movements of the Dolphin-Union caribou herd. Whereas previous authors have examined isolation-by-distance in wolves, this study is the first to demonstrate correlations between genetic structure of wolf populations and the presence of topographical barriers between them. Perhaps most interesting is the possibility that these barriers reflect prey specialization by wolves in different regions.     

Territory size of wolves Canis lupus: linking local (Białowieża Primeval Forest, Poland) and Holarctic-scale patterns

Factors affecting territory size in wolves Canis lupus were studied at 2 scales, the local population (Bia?owie?a Primeval Forest (BPF), eastern Poland) and the geographic range of species (literature review from 14 localities in the Holarctic). Four packs of wolves were studied by radio‐tracking in BPF from 1994 to 1999. The annual territories of packs (Minimum convex polygons with 95% of locations) averaged 201 km² (SD 63, range 116–310). Core areas of territories (50% MCP) covered from 14 to 78 km² (mean 35). Territory sizes and core areas both were negatively correlated to the encounter rates of ungulates (mean number of ungulates seen per unit time spent in the forest by human observers). Pack size (3–8 wolves) did not influence territory size. Home ranges of individual wolves from the same pack varied with season as well as the age, sex, and reproductive status of the wolf. Review of literature from North America and Europe (42–66^oN), showed that latitude and prey biomass were essential factors shaping the biogeographic variation in wolf territory size. Territories increased with latitude and declined with growing biomass of prey. The analysis showed that latitude acted partly independently of the south–north gradient in prey abundance. At similar standing crop of ungulate biomass (100 kg km^?2), wolf territories would average 140 km² at 40^oN, 370 km² at 50^oN, and 950 km² at 60^oN. Pack size was larger at northern latitudes, but the increase did not keep pace with enlargement of territories. Within‐territory density of wolves declined from 2.5–3 wolves 100 km^?2 at 40–45^oN to 0.7 wolves 100 km^?2 at 60^oN. Our analyses documented similarities regarding the role of prey resources in shaping wolf territoriality at the different scales. Furthermore, a macroecological approach revealed additional factors affecting wolf territory size that were not emergent from knowledge of local population.     

Response of Moose Hunters to Predation following Wolf Return in Sweden

Background

Predation and hunter harvest constitute the main mortality factors affecting the size and dynamics of many exploited populations. The re-colonization by wolves (Canis lupus) of the Scandinavian Peninsula may therefore substantially reduce hunter harvest of moose (Alces alces), the main prey of wolves.

Methodology/Principal findings

We examined possible effects of wolf presence on hunter harvest in areas where we had data before and after wolf establishment (n = 25), and in additional areas that had been continuously exposed to wolf predation during at least ten years (n = 43). There was a general reduction in the total number of moose harvested (n = 31,827) during the ten year study period in all areas irrespective of presence of wolves or not. However, the reduction in hunter harvest was stronger within wolf territories compared to control areas without wolves. The reduction in harvest was larger in small (500-800 km²) compared to large (1,200-1,800 km²) wolf territories. In areas with newly established wolf territories moose management appeared to be adaptive with regard to both managers (hunting quotas) and to hunters (actual harvest). In these areas an instant reduction in moose harvest over-compensated the estimated number of moose killed annually by wolves and the composition of the hunted animals changed towards a lower proportion of adult females.

Conclusions/Significance

We show that the re-colonization of wolves may result in an almost instant functional response by another large predator—humans—that reduced the potential for a direct numerical effect on the density of wolves’ main prey, the moose. Because most of the worlds’ habitat that will be available for future colonization by large predators are likely to be strongly influenced by humans, human behavioural responses may constitute a key trait that govern the impact of large predators on their prey.     

Habitat selection by wolvesCanis lupus in the uplands and mountains of southern Poland

Using data from the National Wolf Census, carried out in Poland in 2000–2001, and GIS techniques we analysed habitat selection by wolvesCanis lupus Linnaeus, 1758 in uplands and mountains of southern Poland. We compared ten habitat variables and two parameters related to wolf abundance in 52 circular plots (154 km² each) with recorded wolves and 97 randomly selected plots with no signs of wolf presence. Wolf plots were characterized by higher elevation and closer location to the state border than wolf-free plots. Furthermore, wolf plots had higher forest cover, but smaller number of villages and towns and shorter railways and roads than plots without wolves. The best model explaining wolf distribution included forest cover, number of villages, length of roads and railway lines. Compared to northern Poland, the southern part of the country offers worse habitats for wolves due to significantly denser network of settlements and transportation routes.     

Trophic consequences of terrestrial eutrophication for a threatened ungulate

Changes in primary productivity have the potential to substantially alter food webs, with positive outcomes for some species and negative outcomes for others. Understanding the environmental context and species traits that give rise to these divergent outcomes is a major challenge to the generality of both theoretical and applied ecology. In aquatic systems, nutrient-mediated eutrophication has led to major declines in species diversity, motivating us to seek terrestrial analogues using a large-mammal system across 598 000 km² of the Canadian boreal forest. These forests are undergoing some of the most rapid rates of land-use change on Earth and are home to declining caribou (Rangifer tarandus caribou) populations. Using satellite-derived estimates of primary productivity, coupled with estimates of moose (Alces alces) and wolf (Canis lupus) abundance, we used path analyses to discriminate among hypotheses explaining how habitat alteration can affect caribou population growth. Hypotheses included food limitation, resource dominance by moose over caribou, and apparent competition with predators shared between moose and caribou. Results support apparent competition and yield estimates of wolf densities (1.8 individuals 1000 km⁻²) above which caribou populations decline. Our multi-trophic analysis provides insight into the cascading effects of habitat alteration from forest cutting that destabilize terrestrial predator–prey dynamics. Finally, the path analysis highlights why conservation actions directed at the proximate cause of caribou decline have been more successful in the near term than those directed further along the trophic chain.     

Habitat variables associated with wolf (Canis lupus) distribution and abundance in northern Poland

Based on data collected during the National Wolf Census in 2000–01, we analysed the main habitat factors influencing the distribution and abundance of the wolf, Canis lupus, in northern Poland. The study region forms the western border of the continuous Eastern European range of wolves, although attempts at westward dispersal have been observed. Using Geographic Information System techniques, we measured nine habitat variables and three parameters related to wolf occurrence in 134 circular sample plots (radius 7 km, area 154 km² each). We compared 72 plots where wolves were recorded and 62 plots with no signs of wolf presence. Wolf plots were characterized by significantly higher forest cover, less fragmentation of forests, lower density of villages, towns, motorways, and railways than wolf‐free plots. We found a positive correlation between the sum of wolf observations in plots and forest cover. The number of domestic animals killed by wolves was higher in areas with higher indices of wolf abundance and lower forest area. In multiple regression analysis, four independent variables explained 59% of the variation in wolf distribution and abundance in northern Poland: straight‐line distance to continuous range of wolves in Eastern Europe; forest cover; forest fragmentation; and length of major motorways. We conclude that protection of wolves in Poland (since 1998) may not be an adequate conservation measure, especially because of the increasing density of highways and express motorways. Existing forest corridors should be protected and new ones should be restored to ensure long‐term conservation of wolves and allow range expansion into Western Europe.     

Wolf Depredation on Domestic Animals in the Polish Carpathian Mountains

Abstract As wolves (Canis lupus) recover in Poland, their depredation on domestic animals is increasing, as have conflicts between wolves and farmers. From 1998 to 2004, I investigated spatial and temporal patterns of 591 verified incidents of wolf depredation in the eastern part of the Polish Carpathian Mountains. The wolf population I surveyed covered an estimated range of 4,993 km². Depredation occurred over 1,595 km² of that area. Sheep accounted for 84.8% of domestic animals killed by wolves. Depredation on sheep and number of sheep farms attacked by wolves increased between 1998 and 2004 (r² = 0.61, P = 0.04 and r² = 0.89, P = 0.02, respectively). The number of wolf attacks on sheep farms in a given year were negatively correlated to red deer (Cervus elaphus) population numbers (R² = 0.69, P = 0.02). The amount of depredation caused by each of the 4 monitored packs was best explained by farm density in their territories (R² = 0.59, P = 0.004). Number of attacks recorded on farms was positively correlated to distance from the farm to the pack's den and rendezvous sites (R² = 0.16, P = 0.04). Of depredation recorded in the 4 pack's territories I surveyed, 77% occurred in 4 farms with no or inadequate protection. I concluded that wolf depredation in the studied area is opportunistic. Wolf predation intensity is a function of decreasing abundance of red deer, the density of sheep farms, and proximity of farms to the summer activity centers of wolf packs, and it is facilitated by poor husbandry practices. These results can aid in preventing wolf depredation and provide a foundation for a wolf management plan.     

Modeling Cumulative Effects of Climate and Development on Moose,Wolf, and Caribou Populations

Wildlife models focused solely on a single strong influence (e.g., habitat components, wildlife harvest) are limited in their ability to detect key mechanisms influencing population change. Instead, we propose integrated modeling in the context of cumulative effects assessment using multispecies population dynamics models linked to landscape-climate simulation at large spatial and temporal scales. We developed an integrated landscape and population simulation model using ALCES Online as the model-building platform, and the model accounted for key ecological components and relationships among moose (Alces alces), grey wolves (Canis lupus nubilus), and woodland caribou (Rangifer tarandus caribou) in northern Ontario, Canada. We simulated multiple scenarios over 5 decades (beginning 2020) to explore sensitivity to climate change and land use and assessed effects at multiple scales. The magnitude of effect and the relative importance of key factors (climate change, roads, and habitat) differed depending on the scale of assessment. Across the full extent of the study area (654,311km² [ecozonal scale]), the caribou population declined by 26% largely because of climate change and associated predator-prey response, which led to caribou range recession in the southern part of the study area. At the caribou range scale (108,378 km²), which focused on 2 herds in the northern part of the study area, climate change led to a 10% decline in the population and development led to an additional 7% decline. At the project scale (8,331 km²), which was focused more narrowly on the landscape surrounding 4 proposed mines, the caribou population declined by 29% largely in response to simulated development. Given that observed caribou population dynamics were sensitive to the cumulative effects of climate change, land use, interspecific interactions, and scale, insights from the analysis might not emerge under a less complex model. Our integrated modeling framework provides valuable support for broader regional assessments, including estimation of risk to caribou and Indigenous food security, and for developing and evaluating potential caribou recovery strategies. © 2021 The Authors. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.     

Predator densities and white-tailed deer fawn survival

Predation is the dominant source of mortality for white-tailed deer (Odocoileus virginianus) <6 months old throughout North America. Yet, few white-tailed deer fawn survival studies have occurred in areas with 4 predator species or have considered concurrent densities of deer and predator species. We monitored survival and cause-specific mortality from birth to 6 months for 100 neonatal fawns during 2013–2015 in the Upper Peninsula of Michigan, USA, while simultaneously estimating population densities of deer, American black bear (Ursus americanus), coyote (Canis latrans), bobcat (Lynx rufus), and gray wolf (Canis lupus). We estimated fawn predation risk in response to sex, birth mass, and date of birth. Six-month fawn survival pooled among years was 36%, and fawn mortality risk was not related to birth mass, date of birth, or sex. Estimated mean annual deer and predator densities were 334 fawns/100 km², 25.9 black bear/100 km², 23.8 coyotes/100 km², 3.8 bobcat/100 km², and 2.8 wolves/100 km². Despite lower estimated per-individual kill rates, coyotes and black bears were the leading sources of fawn mortality because they had greater densities relative to bobcats and wolves. Our results indicate that the presence of more predator species in a system is not entirely additive in its effect on fawn survival. © The Wildlife Society, 2019     

Camera-trap studies of maned wolf density in the Cerrado and the Pantanal of Brazil

The maned wolf (Chrysocyon brachyurus) is threatened by large-scale habitat loss, in particular due to conversion to agricultural land. This is the first published study on maned wolf density and the first test of individual identification from camera-trap photographs. We present results from two Brazilian regions: the Cerrado and the Pantanal. Using capture–recapture analysis of camera-trap data, we estimated densities per 100 square kilometers of 3.64 ± 0.77 individuals at the Cerrado site and 1.56 ± 0.77 individuals at the Pantanal site. Parallel radio-telemetry studies at the Pantanal site showed that maned wolves occupied home ranges of 39–58 km² (mean = 50.3 ± 7.67 km²). Our study in the Cerrado took place in a private farm with a mixture of agricultural land and native habitats, representative of the majority of the present-day Cerrado. Whereas many other mammalian species have suffered in the region, our results show that the maned wolf may cope better with this highly fragmented landscape than one might have feared. Finally, the paper briefly compares maned wolf density with density of puma (Puma concolor) in the Pantanal site.     

Conservation of wildlife populations: factoring in incremental disturbance

Progressive anthropogenic disturbance can alter ecosystem organization potentially causing shifts from one stable state to another. This potential for ecosystem shifts must be considered when establishing targets and objectives for conservation. We ask whether a predator–prey system response to incremental anthropogenic disturbance might shift along a disturbance gradient and, if it does, whether any disturbance thresholds are evident for this system. Development of linear corridors in forested areas increases wolf predation effectiveness, while high density of development provides a safe‐haven for their prey. If wolves limit moose population growth, then wolves and moose should respond inversely to land cover disturbance. Using general linear model analysis, we test how the rate of change in moose (Alces alces) density and wolf (Canis lupus) harvest density are influenced by the rate of change in land cover and proportion of land cover disturbed within a 300,000 km² area in the boreal forest of Alberta, Canada. Using logistic regression, we test how the direction of change in moose density is influenced by measures of land cover change. In response to incremental land cover disturbance, moose declines occurred where <43% of land cover was disturbed; in such landscapes, there were high rates of increase in linear disturbance and wolf density increased. By contrast, moose increases occurred where >43% of land cover was disturbed and wolf density declined. Wolves and moose appeared to respond inversely to incremental disturbance with the balance between moose decline and wolf increase shifting at about 43% of land cover disturbed. Conservation decisions require quantification of disturbance rates and their relationships to predator–prey systems because ecosystem responses to anthropogenic disturbance shift across disturbance gradients.     

Correlates of Mortality in an Exploited Wolf Population

Abstract We investigated the influence of habitat use on risk of death from hunting and trapping of 55 radiocollared gray wolves (Canis lupus) from an exploited insular population in Southeast Alaska, USA. We compared mortality rates for resident and nonresident wolves and used Cox proportional hazards regression to relate habitat composition within 100-m circular buffers around radiolocations to risk of death of resident and nonresident wolves. In addition, we included covariates representing distances to roads, logged stands, and lakes and streams in those analyses. We also compiled harvest data from 31 harvest units within the study area to compare densities of roads and distances from human settlements with rates of harvest. During our study 39 wolves died, of which 18 were harvested legally, 16 were killed illegally, and 5 died from natural causes. Legal and illegal harvest accounted for >87% of the mortality of radiocollared resident and nonresident wolves. Mean annual survival was 0.54 (SE = 0.17) for all wolves. Annual survival was 0.65 (SE = 0.17) for resident wolves and 0.34 (SE = 0.17) for nonresidents. Very few (19%) nonresident wolves survived to colonize vacant territories or join existing wolf packs. Roads, muskegs, and distances from lakes and streams were covariates positively associated with death of resident wolves. Clear-cuts were positively associated with risk of death of nonresident wolves. Rate of harvest increased with density of roads; however, road densities >0.9 km/km² had little additional effect on harvest rates. Harvest rates decreased with ocean distances from nearest towns or settlements. Roads clearly increased risk of death for wolves from hunting and trapping and contributed to unsustainable rates of harvest. Wildlife managers should consider effects of roads and other habitat features on harvest of wolves when developing harvest recommendations. They should expect substantial illegal harvest where wolf habitat is accessible to humans. Moreover, high rates of mortality of nonresident wolves exposed to legal and illegal harvest may reduce or delay successful dispersal, potentially affecting linkages between small disjunct wolf populations or population segments. We conclude that a combination of conservative harvest regulations and large roadless reserves likely are the most effective measures for conserving wolves where risks from human-caused mortality are high.     

Can Occupancy–Abundance Models Be Used to Monitor Wolf Abundance?

Estimating the abundance of wild carnivores is of foremost importance for conservation and management. However, given their elusive habits, direct observations of these animals are difficult to obtain, so abundance is more commonly estimated from sign surveys or radio-marked individuals. These methods can be costly and difficult, particularly in large areas with heavy forest cover. As an alternative, recent research has suggested that wolf abundance can be estimated from occupancy–abundance curves derived from “virtual” surveys of simulated wolf track networks. Although potentially more cost-effective, the utility of this approach hinges on its robustness to violations of its assumptions. We assessed the sensitivity of the occupancy–abundance approach to four assumptions: variation in wolf movement rates, changes in pack cohesion, presence of lone wolves, and size of survey units. Our simulations showed that occupancy rates and wolf pack abundances were biased high if track surveys were conducted when wolves made long compared to short movements, wolf packs were moving as multiple hunting units as opposed to a cohesive pack, and lone wolves were moving throughout the surveyed landscape. We also found that larger survey units (400 and 576 km²) were more robust to changes in these factors than smaller survey units (36 and 144 km²). However, occupancy rates derived from large survey units rapidly reached an asymptote at 100% occupancy, suggesting that these large units are inappropriate for areas with moderate to high wolf densities (>15 wolves/1,000 km²). Virtually-derived occupancy–abundance relationships can be a useful method for monitoring wolves and other elusive wildlife if applied within certain constraints, in particular biological knowledge of the surveyed species needs to be incorporated into the design of the occupancy surveys. Further, we suggest that the applicability of this method could be extended by directly incorporating some of its assumptions into the modelling framework.     

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.     

Optimal predator management for mountain sheep conservation depends on the strength of mesopredator release

Large predators often suppress ungulate population growth, but they may also suppress the abundance of smaller predators that prey on neonatal ungulates. Antagonistic interactions among predators may therefore need to be integrated into predator–prey models to effectively manage ungulate–predator systems. We present a modeling framework that examines the net impact of interacting predators on the population growth rate of shared prey, using interactions among wolves Canis lupus, coyotes Canis latrans and Dall sheep Ovis dalli dalli as a case study. Wolf control is currently employed on approximately 16 million ha in Alaska to increase the abundance of ungulates for human harvest. We hypothesized that the positive effects of wolf control on Dall sheep population growth could be counteracted by increased levels of predation by coyotes. Coyotes and Dall sheep adult females (ewes) and lambs were radiocollared in the Alaska Range from 1999–2005 to estimate fecundity, age‐specific survival rates, and causes of mortality in an area without wolf control. We used stage‐structured population models to simulate the net effect of wolf control on Dall sheep population growth (λ). Our models accounted for stage‐specific predation rates by wolves and coyotes, compensatory mortality, and the potential release of coyote populations due to wolf control. Wolves were the main predators of ewes, coyotes were the main predators of lambs, and wolves were the main source of mortality for coyotes. Population models predicted that wolf control could increase sheep λ by 4% per year in the absence of mesopredator release. However, if wolf control released coyote populations, our models predicted that sheep λ could decrease by up to 3% per year. These results highlight the importance of integrating antagonistic interactions among predators into predator–prey models, because the net effect of predator management on shared prey can depend critically on the strength of mesopredator release.     

Differentiation of tundra/taiga and boreal coniferous forest wolves: genetics, coat colour and association with migratory caribou

The grey wolf has one of the largest historic distributions of any terrestrial mammal and can disperse over great distances across imposing topographic barriers. As a result, geographical distance and physical obstacles to dispersal may not be consequential factors in the evolutionary divergence of wolf populations. However, recent studies suggest ecological features can constrain gene flow. We tested whether wolf-prey associations in uninterrupted tundra and forested regions of Canada explained differences in migratory behaviour, genetics, and coat colour of wolves. Satellite-telemetry data demonstrated that tundra wolves (n = 19) migrate annually with caribou (n = 19) from denning areas in the tundra to wintering areas south of the treeline. In contrast, nearby boreal coniferous forest wolves are territorial and associated year round with resident prey. Spatially explicit analysis of 14 autosomal microsatellite loci (n = 404 individuals) found two genetic clusters corresponding to tundra vs. boreal coniferous forest wolves. A sex bias in gene flow was inferred based on higher levels of mtDNA divergence (F(ST) = 0.282, 0.028 and 0.033; P < 0.0001 for mitochondrial, nuclear autosomal and Y-chromosome markers, respectively). Phenotypic differentiation was substantial as 93% of wolves from tundra populations exhibited light colouration whereas only 38% of boreal coniferous forest wolves did (chi(2) = 64.52, P < 0.0001). The sharp boundary representing this discontinuity was the southern limit of the caribou migration. These findings show that substantial genetic and phenotypic differentiation in highly mobile mammals can be caused by prey-habitat specialization rather than distance or topographic barriers. The presence of a distinct wolf ecotype in the tundra of North America highlights the need to preserve migratory populations.