Understanding how genetic drift and inbreeding can shape phenotypic traits and the expression of rare genetic variants can be important for conservation management in small and isolated populations. Following the recolonization of the grey wolf (Canis lupus) on the Scandinavian peninsula, the population has remained small and semi-isolated. The population traces back to seven founders, only and is highly inbred. Inbreeding depression has been demonstrated including effects on reproductive output and several congenital deformities. Some wolves also show anomalous coat color, characterized by hypopigmentation displayed as white tail tips, or even larger areas of white fur. In this study, we utilized four decades of monitoring data, dating back to the recolonization of the Scandinavian peninsula in 1983, to examine the occurrence of expressed anomalous coat color investigating its origin and inheritance, genetic architecture, and the effect of inbreeding. SNP genotyping revealed a distinct haplotype linked to the Melanocyte-inducing transcription factor (MITF) gene that co-segregated with anomalous coat color, suggesting a direct causal effect of this variant. The MITF gene regulates mammal melanocyte development, which in turn affects pigmentation. Our findings demonstrate that the identified gene variant is recessive, which in the homozygous state likely causes the disruption of normal melanocyte development, leading to unpigmented or hypopigmented areas. The origin of this haplotype was traced back to a third founder, reproducing for the first time in 1991. Indeed, the genetic constraints and subsequent inbreeding shaped by few founders, small population size, and semi-isolation over several decades point towards the importance of genetic diversity and facilitated gene flow between populations, but also how such vital immigration can bring about unforeseen side effects if inbreeding continues within the immigrant lineages.

Domestication has profoundly influenced the development of human agriculture, and is characterized by intense artificial selection. Feralization occurs when domestic animals return to the wild. Although feralization does not fully reverse domestication traits, the selective pressures and evolutionary mechanisms driving it remain poorly understood. In northern Europe and Asia, wild, semi-domesticated and feral reindeer (Rangifer tarandus) populations coexist, offering a unique opportunity to study feralization. We studied genomic signatures of selection in semi-domesticated, feral and wild reindeer populations in Norway. Analysis of population structure confirmed historical records indicating the feral population formed a monophyletic clade that originated from neighbouring semi-domesticated reindeer. However, we also found evidence for excess allele-sharing between wild and feral reindeer, indicating hybridization between these lineages. Signatures of selection were identified using a top-down approach (orthologues of domestication candidate genes) and a bottom-up approach (FST and extended haplotypes). We found that candidate regions under selection have higher rates of excess allele-sharing between wild and feral populations. This provides the first evidence of reversed selection pressures during feralization in reindeer and reveals a link between selection and hybridization. Together, our findings suggest that hybridization likely promoted adaptive introgression during feralization. We discuss the implications of our findings for conserving the genetic integrity of endangered wild reindeer populations.

Effective management of the wolf ( Canis lupus ) population in Finland requires defining a genetics-based minimum viable population (MVP) to support the national Favourable Reference Population (FRP) reporting under the EU Habitats Directive. An analysis of long-term and contemporary genetic data from Finnish wolves, together with samples from Scandinavia and Russian Karelia, was carried out to quantify trends in individual heterozygosity, contemporary and historical effective population sizes ( N e ), genetic substructure, and gene flow across regions. Temporal analysis confirmed a persistent decline in genetic diversity and increasing inbreeding levels in Finland over the last two decades. Contemporary data show that the Finnish population consists of two genetically distinct subpopulations; Western Finland and Eastern Finland/Russia, which are connected by low migration rates. These subpopulations are further connected to the Scandinavian and Russian Karelian wolf populations, again with similarly low effective migration rates. Under current demographic conditions and migration rates, neither the Finnish population nor the broader Fennoscandian metapopulation were predicted to reach genetically sustainable long‑term effective population sizes. Applying established short‑term conservation thresholds (the 100/1000 rule) to the two Finnish subpopulations would require a combined MVP of 628 wolves over the next five generations to avoid rising inbreeding levels. Overall, the results demonstrate continuing genetic deterioration in the Finnish wolf population and emphasise that long‑term viability cannot be achieved within Finland alone. Long‑term thresholds ( N e ≥ 1000), corresponding to census sizes >3000 wolves, are not ecologically feasible within Finland. It is therefore vital to ensure connectivity within Fennoscandia, particularly across the Finnish-Russian border. This is essential for preventing inbreeding depression and loss of long-term evolutionary adaptability in a changing environment.

Expanding populations of mesopredators threaten biodiversity and human health in many ecosystems across the world. Lethal control through harvest is commonly implemented as a mitigation measure, yet its effects on mesopredator population dynamics in interaction with compensatory mechanisms and environmental conditions have rarely been assessed quantitatively due to data constraints. Recent advances involving integrated population models (IPMs) have enabled promising new avenues for overcoming these constraints by jointly analyzing multiple datasets while simultaneously accounting for bias and uncertainty. Here we developed a versatile IPM workflow for studying mesopredator population dynamics under different management regimes and applied it to an expanding population of red foxes in Arctic Norway. Our model combined routinely collected data on age, reproductive status, and genetic similarity from >4000 harvested red foxes with opportunistic field observations and information published on red foxes elsewhere. This allowed us to quantify population dynamics over a period of 20 years, and identify the drivers of changes in population growth rates using retrospective (transient Life Table Response Experiments, tLTREs) and prospective (population viability analyses, PVAs) perturbation analyses. We found dramatic year-to-year fluctuations in red fox population size due to natural mortality and immigration responding to changes in rodent prey availability and population density. Forward projections indicated that current harvest levels were likely sufficient to prevent population increase over longer time periods. However, even substantial increases in harvest levels were unable to evoke population decline due to strong buffering effects of density dependence, especially through immigration. Our study highlights the potential of IPMs for studying population dynamics even when no structured surveys of living animals are available, and illustrates the value of extracting and curating information from harvested animals. Our semi-automated and reproducible modeling workflow can be rerun periodically when new data become available for our study population. As the workflow is also designed to be easily adapted for other harvested species, it contributes to the development of cost-effective population analyses that help inform management strategies and mitigate biodiversity loss.

The Scandinavian wolf population (Norway and Sweden) is intensively managed at a population size considered sustainable by managing authorities. As these authorities have decided to reduce the population to only 170 individuals, it is timely to evaluate this population size goal and to scrutinize underlying assumptions of recent simulation studies that inform population management. The effective size of a population determines the pace at which genetic diversity declines and inbreeding increases and plays a crucial role in short- and long-term extinction risks. Here, we use the complete published pedigree of the Scandinavian wolf population to precisely calculate the effective size Ne since the founding of the population, per year. Our results indicate the Ne is unsustainably low. Moreover, we find that recent simulation studies commissioned by the managing authorities greatly overestimate the effective size of the Scandinavian wolf population, questioning their usefulness to inform population management.

Abstract Background The Norwegian lemming (Lemmus lemmus) is a small rodent endemic to the Fennoscandian alpine and arctic tundra. The species is known for cyclic population outbreaks and mass movements during peak years. Previous research based on microsatellites revealed high genetic variation but a weak population structure in the Norwegian lemming. Results In this study, we revisit the population structure of the species using genome-wide data. To do this, we generated a high-quality de novo reference genome for Lemmus lemmus, and resequenced genomes to 2.5–5 × coverage, from 86 lemmings sampled across the species’ entire geographic distribution. Our results reveal that the population is geographically structured into distinct subpopulations, with an overall pattern characterised by isolation-by-distance among subpopulations. Furthermore, our results are consistent with earlier work suggesting that the species survived the last ice age within a northern refugium. Conclusions Together, these findings provide a genome-wide perspective on today’s population structure of the Norwegian lemming. In addition, we provide a de novo reference genome, which we believe will be a valuable resource to the research community.

To reach reproduction, individuals must survive the juvenile stage, a critical period of low survival rates in large carnivores. Early-life conditions during this stage can have lasting effects on survival, reproductive maturation, growth, physiology and behaviour. We assessed recruitment probability in Scandinavian wolves, i.e. the probability that a wolf reaches the reproductive stage and has pups surviving at least five months of age. To unravel human-related and biological factors within the natal territory that could affect recruitment probability, we analysed life-history data from 582 Scandinavian wolves Canis lupus identified by DNA as pups or juveniles in their birth territory. Factors considered included main prey density, road density, human density, and proximity to non-breeding zones, as well as sex, inbreeding level and collaring. Among the 582 wolves analysed, 122 produced at least one surviving pup, corresponding to a recruitment probability of 0.21. Recruitment probability was more than twice as high (0.5) for juvenile wolves fitted with GPS-collars compared to non-collared individuals (0.22), and was positively correlated with human population density in the natal territory. We found no significant effects of other biologically or human-related predictors. These results suggest that in this large carnivore population, managed below carrying capacity, individual recruitment probability is primarily influenced by human-related factors, potentially reflecting poaching risk.

Humans have relied on animal fur for centuries, yet fur farming only began recently during the mid-19th Century. Little is known about this incipient domestication or the genomic processes involved. Domestication may involve founder effects, population bottlenecks and low population size, which, when combined with intense artificial selection, lead to inbreeding, a limited gene pool and reduced fitness. The arctic fox (Vulpes lagopus) has been farmed intensively since the early 1900s and has been artificially selected for economic phenotypes. We investigated the origin of these lineages and the genomic consequences of intensive farming by comparing the genomes of farmed and wild arctic foxes from across their range. Our research indicates recent inbreeding through long Runs of Homozygosity and reduced genomic variation in farmed foxes relative to their respective wild populations. We identified a coastal ecotype origin for all Fennoscandian farmed arctic foxes, aligning them phylogenetically with the wild Icelandic population, a geographically isolated and phenotypically distinct coastal lineage. The depleted genome-wide heterozygosity and increased recent inbreeding in farmed fox lineages is consistent with a heavy consequence of domestication, shedding light on the demographic history and genomic consequences of human manipulation. We highlight the need for increased genomic investigations into fur farm populations to understand the incipient domestication process and uncover the cost of intense farming. The genomic consequences of domestication must be considered in the management of fur farms, with actionable steps needed to prevent descendants of escaped farmed foxes from polluting the gene pool in the wild through introgression.

Demographic declines have important consequences for population viability, since they can lead to losses in genome diversity, as well as increased inbreeding and expression of deleterious mutations. Scandinavia was colonized by the Arctic fox (Vulpes lagopus) at the Pleistocene/Holocene transition, and the population has since been on the periphery of the global distribution. The Scandinavian population became even more fragmented in the early 1900s due to human persecution, and experienced an additional decline in the 1980s. We generated high-coverage genomes from pre-bottleneck, as well as modern Scandinavian and Russian specimens, and found that genome-wide diversity was lower and inbreeding higher in Scandinavia compared to the Siberian population, even prior to the historical bottleneck, most likely reflecting the long-term partial isolation and recent postglacial origin of the Scandinavian population. The southern subpopulation has the highest inbreeding levels, likely due to having been recently founded and highly isolated. Our results also show that although inbreeding increased substantially over the past century, the amount of total genetic load did not change. Overall, these findings illustrate the utility of a temporal approach to disentangle the genomic consequences of recent declines from ancient biogeographic processes.

To reach reproduction, individuals must survive the juvenile stage, a critical period of low survival rates in large carnivores. We analysed data from 582 wolves (Canis lupus) identified by DNA during their first year in Sweden and Norway, to investigate intrinsic and extrinsic factors within the natal territory affecting the probability to reach reproduction, i.e. having pups surviving at least 5 months of age. Factors included main prey density, road density, human density, and proximity to non-breeding zones, as well as sex, inbreeding and being collared. Of the 582 wolves identified, 21% reached reproduction. Human density and whether a wolf was collared were the most significant factors. Both were associated with an increased probability to reach reproduction, potentially linked to poaching. Degree of inbreeding was negatively associated with the probability to reach reproduction, while gravel road density and being born in Sweden were positively associated with it. Our findings suggest an influence of legal and illegal human activities on the juvenile stage of wolves for the probability to reach reproduction. Our study enhances the understanding of how early-life conditions and intrinsic traits shape reproduction and underscores the challenges of wolf conservation in anthropized landscapes. ### Competing Interest Statement The authors have declared no competing interest.

The arctic fox metapopulation in Fennoscandia has been critically endangered over a century, but due to thorough research and intensive conservation measures it has now started to recover. On Varanger Peninsula in northeastern Norway, research about the causes of the arctic fox decline was initiated in 2004. A conceptual model that hypothesized the irregularity of lemming cycles caused by warmer winters and high abundance of the competitively superior red fox as the two main drivers of the critical state of the local population guided the establishment of a monitoring program that included arctic and red foxes, but also other key ecosystem components. Intensive red fox culling in part of the study area was implemented as an experimental management action. This research has subsequently become a part of the Climate-ecological Observatory for Arctic Tundra (COAT), an adaptive monitoring program for long-term research on climate change impacts on terrestrial arctic ecosystems. Following the paradigm of adaptive monitoring, the results of the first 12 years of the study was evaluated in 2017 and led to the conclusion that red fox culling was not sufficient to help arctic foxes recover in the present ecosystem conditions. We hypothesized that absence of lemming peaks and an initially very small and isolated arctic fox population (i.e. demographic stochasticity) contributed to the lack of a response to the culling action. Consequently, two additional management actions decided by the Norwegian Environment Agency were implemented in collaboration with the National Program for Arctic Fox Conservation to address these hypotheses. Supplemental feeding was initiated together with the release of 65 captive bred arctic fox pups from the Norwegian Captive Breeding Program on Varanger over three years (2018-2020). These combined management actions resulted in a strong increase of the local arctic fox population. At the same time, the research conducted to evaluate these management actions has resulted in several now 20-year long time series that are integrated in the research plan of COAT. These data provide unique knowledge about this low arctic ecosystem affected by rapid climate change and increasing human activities. This study shows how on the one hand a monitoring program emerged from a management question – the conservation of arctic foxes – and on the other hand how a thoroughly designed adaptive monitoring program provides crucial information for managing endangered populations.

Wildlife populations are not static. Intrinsic and extrinsic factors affect individuals, which lead to spatiotemporal variation in population density and range. Yet, dynamics in density and their drivers are rarely documented, due in part to the inherent difficulty of studying long-term population-level phenomena at ecologically meaningful scales. We studied the spatiotemporal density dynamics in a recolonizing large carnivore population, the wolverine Gulo gulo , across the Scandinavian Peninsula over nine years. We fitted open-population spatial capture-recapture models to noninvasive genetic sampling data collected across Norway and Sweden to estimate annual density surfaces and their drivers. This approach allowed us to model sex-specific changes in wolverine density and the effect of landscape-level environmental determinants over time. Our results revealed that, as wolverines successfully recolonized many parts of their historical range in Scandinavia, the relationship with spatial determinants of density has changed over time. We also found support for sex-specific responses of the Scandinavian wolverine to the environmental determinants of density and differences in the temporal dynamics of their relationships, indicating disproportionate recolonization ability and anthropogenic pressures. We observed significant changes in the relationship of female wolverine density with several determinants during the study period, suggesting still ongoing expansion of female wolverines whereas males might have already reached the range limits. These findings show that the Scandinavian wolverine population is still recovering from centuries of persecution and severe range contraction. Our study sheds light on the dynamics and challenges of recolonizing large carnivores in human-dominated landscapes across time and space.

Wildlife responses to habitat loss and fragmentation are a central concern in the management and conservation of biodiversity. Small and isolated populations are vulnerable, both due to demographic and genetic mechanisms, which are often linked. Thus, understanding how (changes in) genetic diversity, effective population sizes, and levels of inbreeding relate to population size and degree of isolation is key for developing effective conservation strategies. High-density Single Nucleotide Polymorphism (SNP) arrays represent an increasingly cost-efficient tool to achieve the data needed for such analysis. Here, we present the development of a novel 625k SNP array for reindeer Rangifer tarandus and apply this array to assess conservation genetic issues across thirteen Norwegian wild reindeer populations of varying size, isolation, and genetic origin (i.e., semi-domesticated reindeer origin or a mix of wild reindeer and semi-domesticated reindeer origins). Many of these populations are currently completely isolated, with no gene flow from other populations. We genotyped n = 510 individuals sampled by hunters and found that variation in population size across the populations largely predicted their (recent loss of) genetic variation (observed heterozygosity, Ho), as well as effective population size (Ne) and (change in) level of recent inbreeding. For the smallest and most isolated populations, with total population sizes of <50-100 individuals and a high and increasing level of recent inbreeding, estimated loss of genetic variation was as high as 3-10% over the time span of a generation or less, and estimated Ne was as low as six individuals. With the current level of isolation and associated lack of gene flow, and considering their already low genetic diversity, these populations are hardly viable – neither demographically nor genetically – in the long term. These results have direct relevance for the management of Norwegian wild reindeer, recently red-listed as ‘Near Threatened’. Yet, these genetic challenges, characterizing many of the small ‘wild reindeer’ populations in Norway, have been largely ignored by management thus far. Mitigation efforts such as reducing barriers would introduce substantial conservation dilemma due to the aim of avoiding further spread of chronic wasting disease (CWD), as well as potential further domestic introgression into populations with genetically wild reindeer (or mixed) origin. Nevertheless, our cost-efficient and high-density SNP array especially designed for reindeer and caribou offers a powerful genetic tool to include in future monitoring, providing important contributions to management and conservation decisions. ### Competing Interest Statement The authors have declared no competing interest.

AbstractHarvesting and culling are methods used to monitor and manage wildlife diseases. An important consequence of these practices is a change in the genetic dynamics of affected populations that may threaten their long‐term viability. The effective population size (Ne) is a fundamental parameter for describing such changes as it determines the amount of genetic drift in a population. Here, we estimate Ne of a harvested wild reindeer population in Norway. Then we use simulations to investigate the genetic consequences of management efforts for handling a recent spread of chronic wasting disease, including increased adult male harvest and population decimation. The Ne/N ratio in this population was found to be 0.124 at the end of the study period, compared to 0.239 in the preceding 14 years period. The difference was caused by increased harvest rates with a high proportion of adult males (older than 2.5 years) being shot (15.2% in 2005–2018 and 44.8% in 2021). Increased harvest rates decreased Ne in the simulations, but less sex biased harvest strategies had a lower negative impact. For harvest strategies that yield stable population dynamics, shifting the harvest from calves to adult males and females increased Ne. Population decimation always resulted in decreased genetic variation in the population, with higher loss of heterozygosity and rare alleles with more severe decimation or longer periods of low population size. A very high proportion of males in the harvest had the most severe consequences for the loss of genetic variation. This study clearly shows how the effects of harvest strategies and changes in population size interact to determine the genetic drift of a managed population. The long‐term genetic viability of wildlife populations subject to a disease will also depend on population impacts of the disease and how these interact with management actions.

Harmonising methodology between countries is crucial in transborder population monitoring. However, immediate application of alleged, established DNA-based methods across the extended area can entail drawbacks and may lead to biases. Therefore, genetic methods need to be tested across the whole area before being deployed. Around 4,500 brown bears (Ursus arctos) live in Norway, Sweden, and Finland and they are divided into the western (Scandinavian) and eastern (Karelian) population. Both populations have recovered and are connected via asymmetric migration. DNA-based population monitoring in Norway and Sweden uses the same set of genetic markers. With Finland aiming to implement monitoring, we tested the available SNP-panel developed to assess brown bears in Norway and Sweden, on tissue samples from a representative set of 93 legally harvested individuals from Finland. The aim was to test for ascertainment bias and evaluate its suitability for DNA-based transnational-monitoring covering all three countries. We compared results to the performance of microsatellite genotypes of the same individuals in Finland and against SNP-genotypes from individuals sampled in Sweden (N = 95) and Norway (N = 27). In Finland, a higher resolution for individual identification was obtained for SNPs (PI = 1.18E-27) compared to microsatellites (PI = 4.2E-11). Compared to Norway and Sweden, probability of identity of the SNP-panel was slightly higher and expected heterozygosity lower in Finland indicating ascertainment bias. Yet, our evaluation show that the available SNP-panel outperforms the microsatellite panel currently applied in Norway and Sweden. The SNP-panel represents a powerful tool that could aid improving transnational DNA-based monitoring of brown bears across these three countries.

Conservation reintroduction is an increasingly applied tool for population restoration. The choice of source population is important from a conservation genetic perspective, but there is ambiguity in IUCN policies surrounding reintroductions from (semi-)domestic sources. After the depopulation of an entire wild reindeer population due to detection of chronic wasting disease in Norway, the plan is to re-establish reindeer after years of fallowing. Establishing wild reindeer has become challenging due to arguments for disease control favoring a semidomestic origin of the reintroduced stock. Selection is strong during domestication, and feralization does not lead to a full reversal towards wild traits. From a conservation perspective, we advocate future guidelines to include a ranking when wild, captive, semidomestic, and domestic stocks are available for reintroductions, in order to avoid feralization whenever possible during reintroductions and rewilding efforts.

Abstract Thiamine deficiency can result in life‐threatening physiological and neurological complications. While a thiamine‐deficient diet may result in the onset of such symptoms, the presence of thiaminase – an enzyme that breaks down thiamine – is very often the cause. In such instances, thiaminase counteracts the bioavailability and uptake of thiamine, even when food‐thiamine levels are adequate. Here, we report on a case of failed reproduction in seven Arctic fox (Vulpes lagopus) breeding pairs kept at a captive breeding facility, including the presentation of severe thiamine deficiency symptoms in two male foxes. Symptoms included ataxia, obtundation, truncal sway, star‐gazing and visual impairment. Blood tests were inconclusive, yet symptoms resolved following treatment with a series of thiamine hydrochloride injections, thereby verifying the diagnosis. A fish‐dominated feed, which for the first time had been frozen for a prolonged period, was identified as the likely source of thiaminase and subsequent deterioration in the animals’ health. Symptoms in the two males arose during the annual mating period. All seven breeding pairs at the captive breeding station failed to reproduce – a phenomenon never recorded during the captive breeding facility's preceding 17‐year operation. Relating our findings to peer‐reviewed literature, the second part of this case report assesses how thiamine deficiency (due to thiaminase activity) likely resulted in subclinical effects that impaired the production of reproduction hormones, and thereby led to a complete breeding failure. While previous work has highlighted the potentially lethal effects of thiamine deficiency in farmed foxes, this is, to our knowledge the first study showing how subclinical effects in both males and females may inhibit reproduction in foxes in general, but specifically Arctic foxes. The findings from our case report are not only relevant for captive breeding facilities, but for the welfare and management of captive carnivorous animals in general.

Dedicated conservation efforts spanning the past two decades have saved the Fennoscandian Arctic fox (Vulpes lagopus) population from local extinction, and extensive resources continue to be invested in the species' conservation and management. Although increasing, populations remain isolated, small and are not yet viable in the longer term. An understanding of causes of mortality are consequently important to optimize ongoing conservation actions. Golden eagles (Aquila chrysaetos) are a predator of Arctic foxes, yet little information on this interaction is available in the literature. We document and detail six confirmed cases of Golden eagle depredation of Arctic foxes at the Norwegian captive breeding facility (2019-2022), where foxes are housed in large open-air enclosures in the species' natural habitat. Here, timely detection of missing/dead foxes was challenging, and new insights have been gained following recently improved enclosure monitoring. Golden eagle predation peaked during the winter months, with no cases reported from June to November. This finding contrasts with that which is reported from the field, both for Arctic and other fox species, where eagle depredation peaked at dens with young (summer). While the seasonality of depredation may be ecosystem specific, documented cases from the field may be biased by higher survey efforts associated with the monitoring of reproductive success during the summer. Both white and blue color morphs were housed at the breeding station, yet only white foxes were preyed upon, and mortality was male biased. Mitigation measures and their effectiveness implemented at the facility are presented. Findings are discussed in the broader Arctic fox population ecology and conservation context.

Scavenging is an important part of food acquisition for many carnivore species that switch between scavenging and predation. In landscapes with anthropogenic impact, humans provide food that scavenging species can utilize. We quantified the magnitude of killing versus scavenging by gray wolves (Canis lupus) in Scandinavia where humans impact the ecosystem through hunter harvest, land use practices, and infrastructure. We investigated the cause of death of different animals utilized by wolves, and examined how the proportion of their consumption time spent scavenging was influenced by season, wolf social affiliation, level of inbreeding, density of moose (Alces alces) as their main prey, density of brown bear (Ursus arctos) as an intraguild competitor, and human density. We used data from 39 GPS-collared wolves covering 3198 study days (2001-2019), including 14,205 feeding locations within space-time clusters, and 1362 carcasses utilized by wolves. Most carcasses were wolf-killed (80.5%) while a small part had died from other natural causes (1.9%). The remaining had either anthropogenic mortality causes (4.7%), or the cause of death was unknown (12.9%). Time spent scavenging was higher during winter than during summer and autumn. Solitary wolves spent more time scavenging than pack-living individuals, likely because individual hunting success is lower than pack success. Scavenging time increased with the mean inbreeding coefficient of the adult wolves, possibly indicating that more inbred individuals resort to scavenging, which requires less body strength. There was weak evidence for competition between wolves and brown bears as well as a positive relationship between human density and time spent scavenging. This study shows how both intrinsic and extrinsic factors drive wolf scavenging behavior, and that despite a high level of inbreeding and access to carrion of anthropogenic origin, wolves mainly utilized their own kills.