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1. Estimating the contribution of demographic parameters to changes in population growth is essential for understanding why populations fluctuate. Integrated population models (IPMs) offer a possibility to estimate the contributions of additional demographic parameters, for which no data have been explicitly collected— typically immigration. Such parameters are often subsequently highlighted as important drivers of population growth. Yet, accuracy in estimating their temporal variation, and consequently their contribution to changes in population growth rate, has not been investigated. 2. To quantify the magnitude and cause of potential biases when estimating the contribution of immigration using IPMs, we simulated data (using northern wheatear Oenanthe oenanthe population estimates) from controlled scenarios to examine potential biases and how they depend on IPM parameterization, formulation of priors, the level of temporal variation in immigration and sample size. We also used empirical data on populations with known rates of immigration: Soay sheep Ovis aries and Mauritius kestrel Falco punctatus with zero immigration and grey wolf Canis lupus in Scandinavia with near-zero immigration. 3. IPMs strongly overestimated the contribution of immigration to changes in population growth in scenarios when immigration was simulated with zero temporal variation (proportion of variance attributed to immigration = 63% for the more constrained formulation and real sample size) and in the wild populations, where the true number of immigrants was zero or near-zero (kestrel 19.1%–98.2%, sheep 4.2%– 36.1% and wolf 84.0%–99.2%). Although the estimation of the contribution of immigration in the simulation study became more accurate with increasing temporal variation and sample size, it was often not possible to distinguish between an accurate estimation from data with high temporal variation versus an overestimation from data with low temporal variation. Unrealistically, large sample sizes may be required to estimate the contribution of immigration well. 4. To minimize the risk of overestimating the contribution of immigration (or any additional parameter) in IPMs, we recommend to: (a) look for evidence of variation in immigration before investigating its contribution to population growth, (b) simulate and model data for comparison to the real data and (c) use explicit data on immigration when possible.

Significance We are experiencing the accelerated loss and reconfiguration of biological diversity. Meanwhile, those charged with natural resource management are struggling to meet the challenges of monitoring and managing wildlife populations across vast areas crisscrossed by political borders. What if, akin to weather maps, we could track and forecast the dynamics of wildlife populations across space and time? Using the world’s most extensive large carnivore monitoring program, we showcase the application of an effective tool for spatially explicit quantification of wildlife population dynamics at scales that are relevant to management and conservation., The ongoing recovery of terrestrial large carnivores in North America and Europe is accompanied by intense controversy. On the one hand, reestablishment of large carnivores entails a recovery of their most important ecological role, predation. On the other hand, societies are struggling to relearn how to live with apex predators that kill livestock, compete for game species, and occasionally injure or kill people. Those responsible for managing these species and mitigating conflict often lack fundamental information due to a long-standing challenge in ecology: How do we draw robust population-level inferences for elusive animals spread over immense areas? Here we showcase the application of an effective tool for spatially explicit tracking and forecasting of wildlife population dynamics at scales that are relevant to management and conservation. We analyzed the world’s largest dataset on carnivores comprising more than 35,000 noninvasively obtained DNA samples from over 6,000 individual brown bears (Ursus arctos), gray wolves (Canis lupus), and wolverines (Gulo gulo). Our analyses took into account that not all individuals are detected and, even if detected, their fates are not always known. We show unequivocal quantitative evidence of large carnivore recovery in northern Europe, juxtaposed with the finding that humans are the single-most important factor driving the dynamics of these apex predators. We present maps and forecasts of the spatiotemporal dynamics of large carnivore populations, transcending national boundaries and management regimes.

Swayne's hartebeest Alcelaphus buselaphus swaynei was once widely distributed in the Horn of Africa. By the early 20th century, however, it was extirpated across most of its range and is now limited to two relict populations in the Ethiopian Rift Valley and categorized as Endangered on the IUCN Red List. In this study, we estimated the size and genetic diversity of these two remaining populations, with a particular focus on competition with livestock. We used a total block count method for both Swayne's hartebeest and livestock population counts, and faecal samples for a population genetic analysis. We estimated the total population of Swayne's hartebeest to be 1,528, with 518 individuals in Senkele Swayne's Hartebeest Sanctuary and 1,010 individuals in Maze National Park. Livestock densities were 212 and 153 times those of Swayne's hartebeest in Senkele Swayne's Hartebeest Sanctuary and Maze National Park, respectively. Among 73 mitochondrial D-loop sequences (34 from Senkele Swayne's Hartebeest Sanctuary and 39 from Maze National Park), we found 22 haplotypes (Senkele 12, Maze 16, shared 6). Population genetic parameters suggest only weak sub-structuring between the two populations (FST = 0.164). Despite the positive population trends in both protected areas, the spatial overlap with livestock may lead to future population decline as a result of resource competition and disease transmission. We therefore recommend further translocation to other protected areas within the species’ former range.

Populations of large mammals have declined at alarming rates, especially in areas with intensified land use where species can only persist in small habitat fragments. To support conservation planning, we developed habitat suitability models for the Walia ibex (Capra walie), an endangered wild goat endemic to the Simen Mountains, Ethiopia. We calibrated several models that differ in statistical properties to estimate the spatial extent of suitable habitats of the Walia ibex in the Simen Mountains, as well as in other parts of the Ethiopian highlands to assess potentially suitable areas outside the current distribution range of the species. We further addressed the potential consequences of future climate change using a climate model with four emission scenarios. Model projections estimated the potential suitable habitat under current climate to 501–672 km2 in Simen and 6,251–7,732 km2 in other Ethiopian mountains. Under projected climate change by 2,080, the suitable habitat became larger in Simen but smaller in other parts of Ethiopia. The projected expansion in Simen is contrary to the general expectation of shrinking suitable habitats for high-elevation species under climate warming and may partly be due to the ruggedness of these particular mountains. The Walia ibex has a wide altitudinal range and is able to exploit very steep slopes, allowing it to track the expected vegetation shift to higher altitudes. However, this potential positive impact may not last long under continued climate warming, as the species will not have much more new space left to colonize. Our study indicates that the current distribution range can be substantially increased by reintroducing and/or translocating the species to other areas with suitable habitat. Indeed, to increase the viability and prospects for survival of this flagship species, we strongly recommend human-assisted reintroduction to other Ethiopian mountains. Emulating the successful reintroduction of the Alpine ibex that has spread from a single mountain in Italy to its historical ranges of the Alps in Europe might contribute to saving the Walia ibex from extinction.

1. Selection for crypsis has been recognized as an important ecological driver of animal colouration, whereas the relative importance of thermoregulation is more contentious with mixed empirical support. A potential thermal advantage of darker individuals has been observed in a wide range of animal species. Arctic animals that exhibit colour polymorphisms and undergo seasonal colour moults are interesting study subjects for testing the two alternative hypotheses: demographic performance of different colour morphs might be differentially affected by snow cover with a cryptic advantage for lighter morphs, or conversely by winter temperature with a thermal advantage for darker morphs. 2. In this study, we explored whether camouflage and thermoregulation might explain differences in reproduction and survival between the white and blue colour morphs of the Arctic fox Vulpes lagopus under natural conditions. 3. Juvenile and adult survival, breeding propensity and litter size were measured for 798 captive-bred and released or wild-born Arctic foxes monitored during an 11-year period (2007–2017) in two subpopulations in south-central Norway. We investigated the proportion of the two colour morphs and compared their demographic performance in relation to spatial variation in duration of snow cover, onset of snow season and winter temperatures. 4. After population re-establishment, a higher proportion of blue individuals was observed among wild-born Arctic foxes compared to the proportion of blue foxes released from the captive population. Our field study provides the first evidence for an effect of colour morph on the reproductive performance of Arctic foxes under natural conditions, with a higher breeding propensity of the blue morph compared to the white one. Performance of the two colour morphs was not differentially affected by the climatic variables, except for juvenile survival. Blue morph juveniles showed a tendency for higher survival under colder winter temperatures but lower survival under warmer temperatures compared to white morph juveniles. 5. Overall, our findings do not consistently support predictions of the camouflage or the thermoregulation hypotheses. The higher success of blue foxes suggests an advantage of the dark morph not directly related to disruptive selection by crypsis or thermoregulation. Our results rather point to physiological adaptations and behavioural traits not necessarily connected to thermoregulation, such as stress response, immune function, sexual behaviour and aggressiveness. Our findings highlight the need to explore the potential role of genetic linkage or pleiotropy in influencing the fitness of white and blue Arctic foxes as well as other species with colour polymorphisms. apparent survival, Arctic fox, camouflage, capture–mark– recapture models, colour polymorphism, fitness, reproductive performance, snow cover This is an open access article under the terms of the Creative Commons Attribution-NonCommercial- NoDerivs License, which permits use and distribution in any medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made. © 2021 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society

Spatial capture–recapture modelling (SCR) is a powerful tool for estimating density, population size, and space use of elusive animals. Here, we applied SCR modelling to non-invasive genetic sampling (NGS) data to estimate red fox (Vulpes vulpes) densities in two areas of boreal forest in central (2016–2018) and southern Norway (2017–2018). Estimated densities were overall lower in the central study area (mean = 0.04 foxes per km2 in 2016, 0.10 in 2017, and 0.06 in 2018) compared to the southern study area (0.16 in 2017 and 0.09 in 2018). We found a positive effect of forest cover on density in the central, but not the southern study area. The absence of an effect in the southern area may reflect a paucity of evidence caused by low variation in forest cover. Estimated mean home-range size in the central study area was 45 km2 [95%CI 34–60] for females and 88 km2 [69–113] for males. Mean home-range sizes were smaller in the southern study area (26 km2 [16–42] for females and 56 km2 [35–91] for males). In both study areas, detection probability was session-dependent and affected by sampling effort. This study highlights how SCR modelling in combination with NGS can be used to efficiently monitor red fox populations, and simultaneously incorporate ecological factors and estimate their effects on population density and space use.

Knowledge about the connectivity among natural populations is essential to identify management units for effective conservation actions. Conservation-minded management has led to the recovery of large carnivore populations in northern Europe, possibly restoring connectivity between the two separated, but expanding brown bear (Ursus arctos) populations on the Scandinavian peninsula to the west and Karelia, a part of the large Eurasian population, to the east. The degree of connectivity between these populations has been poorly understood, therefore we investigated the extent of connectivity between the two populations using autosomal microsatellites and Y chromosome haplotypes in 924 male bears (the dispersing sex), sampled during a period of 12 years (2005–2017) across the transborder area where these two populations meet. Our results showed that the two populations are not genetically isolated as reported in earlier studies. We detected recent asymmetrical gene flow at a rate (individuals per generation) of 4.6–5.5 (1%) from Karelia into Scandinavia, whereas the rate was approximately 27.1–34.5 (8%) in the opposite direction. We estimated historical gene flow of effective number of migrants to be between 1.7 and 2.5 between the populations. Analyses of Y chromosome markers supported these results. Successful recovery and expansion of both populations led to the restoration of connectivity, however, it is asymmetric, possibly due to different recovery histories and population densities. By aligning monitoring between neighboring countries, we were able to better understand the biological processes across the relevant spatial scale. Brown bear Genetic structure Male gene flow Microsatellites Migration Recovery Ursus arctos Wildlife monitoring Y chromosome

Genome-wide association studies provide good opportunities for studying the genetic basis of adaptive traits in wild populations. Yet, previous studies often failed to identify major effect genes. In this study, we used high-density single nucleotide polymorphism and individual fitness data from a wild non-model species. Using a whole-genome approach, we identified the MC1R gene as the sole causal gene underlying Arctic fox Vulpes lagopus fur colour. Further, we showed the adaptive importance of fur colour genotypes through measures of fitness that link ecological and evolutionary processes. We found a tendency for blue foxes that are heterozygous at the fur colour locus to have higher fitness than homozygous white foxes. The effect of genotype on fitness was independent of winter duration but varied with prey availability, with the strongest effect in years of increasing rodent populations. MC1R is located in a genomic region with high gene density, and we discuss the potential for indirect selection through linkage and pleiotropy. Our study shows that whole-genome analyses can be successfully applied to wild species and identify major effect genes underlying adaptive traits. Furthermore, we show how this approach can be used to identify knowledge gaps in our understanding of interactions between ecology and evolution.

After decades, even centuries of persecution, large carnivore populations are widely recovering in Europe. Considering the recent recovery of the wolverine (Gulo gulo) in Finland, our aim was to evaluate genetic variation using 14 microsatellites and mtDNA control region (579 bp) in order (1) to determine whether the species is represented by a single genetic population within Finland, (2) to quantify the genetic diversity, and (3) to estimate the effective population size. We found two major genetic clusters divided between eastern and northern Finland based on microsatellites (FST= 0.100) but also a significant pattern of isolation by distance. Wolverines in western Finland had a genetic signature similar to the northern cluster, which can be explained by former translocations of wolverines from northern to western Finland. For both main clusters, most estimates of the effective population sizeNewere below 50. Nevertheless, the genetic diversity was higher in the eastern cluster (HE= 0.57,AR= 4.0,AP= 0.3) than in the northern cluster (HE= 0.49,AR= 3.7,AP= 0.1). Migration between the clusters was low. Two mtDNA haplotypes were found: one common and identical to Scandinavian wolverines; the other rare and not previously detected. The rare haplotype was more prominent in the eastern genetic cluster. Combining all available data, we infer that the genetic population structure within Finland is shaped by a recent bottleneck, isolation by distance, human-aided translocations and postglacial recolonization routes.

The Arctic fox (Vulpes lagopus) population in Fennoscandia experienced a drastic bottleneck in the late 19th century as a result of high hunting pressure. In the 1990s, despite nearly 70 years of protection, the population showed no signs of recovery. In order to mitigate the population decline and facilitate re-establishment, conservation actions including supplementary feeding and red fox culling were implemented in 1998, followed by the reintroduction of foxes from a captive breeding programme, starting in 2006. A positive demographic impact of these actions is evident from a doubling of the population size over the past decade. We used genetic data collected in eight subpopulations between 2008 and 2015 to address whether the recent demographic recovery has been complemented by changes in genetic variation and connectivity between subpopulations. Our results show that genetic variation within subpopulations has increased considerably during the last decade, while genetic differentiation among subpopulations has decreased. A marked shift in metapopulation dynamics is evident during the study period, suggesting substantially increased migration across the metapopulation. This shift followed the recolonization of an extinct subpopulation through the release of foxes from the captive breeding programme and was synchronized in time with the implementation of supplementary feeding and red fox culling in stepping stone patches between core subpopulations in mid-Scandinavia. Indeed, the increase in genetic variation and connectivity in the Scandinavian arctic fox population suggests that metapopulation dynamics have been restored, which may indicate an increase in the long-term viability of the population. This is an open access article distributed under the terms of the Creative Commons CC-BY license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

Estimating wildlife population size is fundamental for wildlife management and conservation. However, making monitoring of population size less resource demanding while still keeping a high monitoring accuracy and precision remains a recurrent challenge. One proposed alternative to count individuals is to instead focus on counting a segment of the population that is easier to monitor but at the same time well informative on total population size. We show how total population size can be estimated from group counts by using an individual-based population model in a social living species. We developed a wolf ( Canis lupus ) specific Individual Based Model and used Approximate Bayesian Computation (ABC) to fit this population model to the time series of annual number of packs, reproductions and pairs obtained from Scandinavian monitoring data. Model informative priors were obtained with data from collared individuals by the Scandinavian wolf research project. The fitted model was then used to estimate a conversion factor from number of packs to total number of individuals and to number of reproductions. There was a good fit between the retained simulations by ABC and the observed Scandinavian wolf population trajectory. The fitted simulations returned a conversion factor of 8.0 (95% CI = 6.62–10.07) from number of packs to total population size and of 1.0 (95% CI = 0.93–1.12) to number of reproductions in December. A sensitivity analysis revealed that the conversion factor from packs to total population size was positively correlated with pup survival and litter size and negatively correlated with subadult, vagrant and adult survivals. Using an individual based model allowed us to model the full complexity of demographic traits of a social-living species such as the wolf. The flexibility of the model also meant that the conversion factor could be estimated for any month during the year. Our approach to estimate total population size from counts of groups requires having a population model where both individuals and groups are explicitly described and can be applied to other wolf populations and group-living species where counting all individuals over a large area is unfeasible.

For socially monogamous species, breeder bond dissolution has important consequences for population dynamics, but the extent to which extrinsic or intrinsic population factors causes pair dissolution remain poorly understood, especially among carnivores. Using an extensive life-history data set, a survival analysis and competing risks framework, we examined the fate of 153 different wolf (Canis lupus) pairs in the recolonizing Scandinavian wolf population, during 14 winters of snow tracking and DNA monitoring. Wolf pair dissolution was generally linked to a mortality event and was strongly affected by extrinsic (i.e. anthropogenic) causes. No divorce was observed, and among the pair dissolution where causes have been identified, death of one or both wolves was always involved. Median time from pair formation to pair dissolution was three consecutive winters (i.e. approximately 2 years). Pair dissolution was mostly human-related, primarily caused by legal control actions (36·7%), verified poaching (9·2%) and traffic-related causes (2·1%). Intrinsic factors, such as disease and age, accounted for only 7·7% of pair dissolutions. The remaining 44·3% of dissolution events were from unknown causes, but we argue that a large portion could be explained by an additional source of human-caused mortality, cryptic poaching. Extrinsic population factors, such as variables describing the geographical location of the pair, had a stronger effect on risk of pair dissolution compared to anthropogenic landscape characteristics. Population intrinsic factors, such as the inbreeding coefficient of the male pair member, had a negative effect on pair bond duration. The mechanism behind this result remains unknown, but might be explained by lower survival of inbred males or more complex inbreeding effects mediated by behaviour. Our study provides quantitative estimates of breeder bond duration in a social carnivore and highlights the effect of extrinsic (i.e. anthropogenic) and intrinsic factors (i.e. inbreeding) involved in wolf pair bond duration. Unlike the effects of intrinsic and extrinsic factors that are commonly reported on individual survival or population growth, here we provide quantitative estimates of their potential effect on the social unit of the population, the wolf pair.

Population expansions of boreal species are among the most substantial ecological consequences of climate change, potentially transforming both structure and processes of northern ecosystems. Despite their importance, little is known about expansion dynamics of boreal species. Red foxes (Vulpes vulpes) are forecasted to become a keystone species in northern Europe, a process stemming from population expansions that began in the 19th century. To identify the relative roles of geographic and demographic factors and the sources of northern European red fox population expansion, we genotyped 21 microsatellite loci in modern and historical (1835-1941) Fennoscandian red foxes. Using Bayesian clustering and Bayesian inference of migration rates, we identified high connectivity and asymmetric migration rates across the region, consistent with source-sink dynamics, whereby more recently colonized sampling regions received immigrants from multiple sources. There were no clear clines in allele frequency or genetic diversity as would be expected from a unidirectional range expansion from south to north. Instead, migration inferences, demographic models and comparison to historical red fox genotypes suggested that the population expansion of the red fox is a consequence of dispersal from multiple sources, as well as in situ demographic growth. Together, these findings provide a rare glimpse into the anatomy of a boreal range expansion and enable informed predictions about future changes in boreal communities.

Understanding the response of boreal species to past climate warming can help to predict future responses to climate change. In the Northern Hemisphere, the distribution and abundance of northern populations have been influenced by previous glaciations. In this study, we investigated the population history of the Fennoscandian red fox (Vulpes vulpes), which is a generalist carnivore currently undergoing range expansion in the tundra ecosystem. By analysing a 696 bp sequence of the mitochondrial DNA (N = 259) and two Y chromosome-specific microsatellite loci (N = 120), we specifically investigated where the red fox survived the Last Glacial Maximum and how Fennoscandia was recolonized. There was high genetic continuity across most of Fennoscandia, and we identified at least two recolonization pathways: one from continental Europe and one from the northeast (Siberia). Mitochondrial haplotype diversity displayed a significant decline with increasing latitude, consistent with expectations of unidirectional colonization. Each region displayed signatures of recent demographic and/or range expansions. For Finland, an additional recolonization route was suggested from the mismatch distribution analysis and identification of novel haplotypes. We concluded that, as with many boreal generalist species, the Fennoscandian red fox originates from multiple refugia, suggesting that it has benefited from diverse evolutionary histories, potentially enhancing its tolerance to different habitat conditions.

Source at http://dx.doi.org/10.1080/17518369.2017.1319602 The biodiversity working group of the Arctic Council has developed pan-Arctic biodiversity monitoring plans to improve our ability to detect, understand and report on long-term change in Arctic biodiversity. The Arctic fox (Vulpes lagopus) was identified as a target of future monitoring because of its circumpolar distribution, ecological importance and reliance on Arctic ecosystems. We provide the first exhaustive survey of contemporary Arctic fox monitoring programmes, describing 34 projects located in eight countries. Monitored populations covered equally the four climate zones of the species’ distribution, and there were large differences between populations in long-term trends, multi-annual fluctuations, diet composition, degree of competition with red fox and human interferences. Den density, number of active dens, number of breeding dens and litter size were assessed in almost all populations, while projects varied greatly with respect to monitoring of other variables indicative of population status, ecosystem state or ecosystem function. We review the benefits, opportunities and challenges to increased integration of monitoring projects. We argue that better harmonizing protocols of data collection and data management would allow new questions to be addressed while adding tremendous value to individual projects. However, despite many opportunities, challenges remain. We offer six recommendations that represent decisive progress toward a better integration of Arctic fox monitoring projects. Further, our work serves as a template that can be used to integrate monitoring efforts of other species, thereby providing a key step for future assessments of global biodiversity.

Inbreeding (mating between relatives) is a major concern for conservation as it decreases the fitness of offspring and can increase the extinction risk of populations. While pedigrees have traditionally been used to measure individual inbreeding, molecular markers have opened up new avenues to characterize inbreeding. However, a limitation has been that small numbers of markers can only roughly measure the proportion of an individual’s genome that is identical-by-descent (IBD) due to inbreeding. We used whole-genome resequencing of 97 grey wolves (Canis lupus) from the highly inbred Scandinavian wolf population, originally founded by only two individuals, to identify IBD chromosome segments as runs of homozygosity (ROH). This gave the very high resolution required to precisely measure the realized IBD fraction of the genome as FROH. We found a striking pattern of complete or near-complete homozygosity of entire chromosomes in many individuals. The majority of individual IBD was contributed by long IBD segments (>5cM) originating from common ancestors of parents within the last ~10 generations. However, although most IBD segments were very short (

Introgression and admixture from domestic or foreign conspecifics into wild populations are of great concern in wildlife conservation. The issue is of particular interest in ungulates where translocations and re-introductions have been common practice. In Europe, the only large remaining wild populations of reindeer (Rangifer tarandus L.) are found in the mountainous habitats of southern Norway. These populations have during the last centuries been exposed to extensive habitat modifications and periods of contact with domestic reindeer. Through analyses of ancient and extant mitochondrial DNA (mtDNA) we document extensive intra- and inter-population genetic changes during the last millennium. Our data indicate population reduction within a short time interval during the 11th–12th century during a period when mass trapping of reindeer was common. Significant differentiation between the ancient herds suggests an ancient genetic structuring of the reindeer herds in southern Norway, although not as strong as between modern herds. Two different mtDNA lineages characterized the ancient herds. A Bayesian approach to reconstruct the recent evolutionary history suggests that the reindeer herds in southern Norway originate from two populations separated in different refugia during the last glacial period. The presence of two additional extant lineages, characteristic of domestic herds, suggests substantial introgression into the native wild reindeer. The putatively different refugial origins of the lineages represented by the extant herds in Rondane/Dovre, Hardangervidda and those with a mainly domestic origin may well reflect different adaptations to environmental conditions, including degree of human interference. Further research on this issue would provide important insights for conservation priorities and a sustainable and flexible management strategy for the remaining wild reindeer herds in the Scandinavian mountains.

Source at http://doi.org/10.1080/17518369.2017.1323621 The distribution of traditional breeding dens on the Varanger Peninsula (70–71°N) in northernmost Fennoscandia indicates that this area once harboured a large Arctic fox population. Early 20th century naturalists regarded the coastal tundra of the Fennoscandian Low Arctic to be a stronghold for the species. At the start of our research in 2004, however, the local Arctic fox population was critically small and most neighbouring populations had been extirpated. Here, we synthesize the results of 11 years of research to highlight ecosystem drivers behind the critical state of the Arctic fox in Low-Arctic Fennoscandia. We identify two fundamental drivers: (1) an increasingly climate-driven irregularity of the lemming cycle and (2) a management- and climate-driven increase in the abundance of red fox that is subsidized by more ungulate carrion. Arctic fox reproductive success is low when lemmings are scarce (despite high vole abundance), while red foxes exclude Arctic foxes from high-quality breeding territories in summer and from marine and terrestrial carrion in winter. Red fox culling on Varanger Peninsula may have prevented the extirpation of the Arctic fox population. However, one decade after the onset of this management action the Arctic fox population has failed to increase either because the action has been insufficient or because demographic and environmental stochasticity has precluded a positive response. We discuss options for future research and management of the Arctic fox in the Fennoscandian Low Arctic.

The Arctic fox (Vulpes lagopus L.) is listed as extinct in Finland, endangered in Sweden and critically endangered in Norway. Around 2000 there were only 40–60 adult individuals left, prompting the implementation of conservation actions, including a captive breeding programme founded from wild-caught pups. The initial breeding trials failed, probably because of stress among captive animals, and the programme was radically changed in 2005. Eight large enclosures within the species’ historical natural habitat were established, which had the positive effect of all pairs breeding in 2007. As of 2015, 385 pups (yearly average 37) were produced. In this ongoing programme, pups are released the winter (January–February) following their birth and have had an average first-year survival of 0.44. The release sites are prepared with artificial dens and a network of supplementary food dispensers, designed to work exclusively for the Arctic fox. After just four to seven years of releases, populations have been effectively re-established in three mountain areas where the species had been locally extinct. One of the newly re-established populations has become the largest population in Norway. Several other populations, including Swedish ones, have benefited considerably from successful immigration of released foxes. The number of wild-born pups that are descendants of released foxes has likely exceeded 600, and in 2014 50% of all free-living breeding pairs in mainland Norway included released foxes or their descendants. The Norwegian Arctic fox captive breeding programme has proven to be an important conservation action for the recovery of the Scandinavian Arctic fox population.

Isolation of small populations can reduce fitness through inbreeding depression and impede population growth. Outcrossing with only a few unrelated individuals can increase demographic and genetic viability substantially, but few studies have documented such genetic rescue in natural mammal populations. We investigate the effects of immigration in a subpopulation of the endangered Scandinavian arctic fox (Vulpes lagopus), founded by six individuals and isolated for 9 years at an extremely small population size. Based on a long-term pedigree (105 litters, 543 individuals) combined with individual fitness traits, we found evidence for genetic rescue. Natural immigration and gene flow of three outbred males in 2010 resulted in a reduction in population average inbreeding coefficient (f), from 0.14 to 0.08 within 5 years. Genetic rescue was further supported by 1.9 times higher juvenile survival and 1.3 times higher breeding success in immigrant first-generation offspring compared with inbred offspring. Five years after immigration, the population had more than doubled in size and allelic richness increased by 41%. This is one of few studies that has documented genetic rescue in a natural mammal population suffering from inbreeding depression and contributes to a growing body of data demonstrating the vital connection between genetics and individual fitness.

Noninvasive genetic monitoring has the potential to estimate vital rates essential for conservation and management of many species. In a long-term genetic capture-mark-recapture study using scats we evaluated temporal variation in adult survival in a wolverine (Gulo gulo) population in southern Norway. In contrast to most previous studies of large mammals we found evidence for negative density dependence in adult survival in this large carnivore. Both sexes showed the same pattern of density dependence, with higher annual survival rates in adult females than males. In addition, we also found an additive mortality effect of harvesting in the population, resulting in the lowest adult survival rates at a combination of high population density and high harvest rate. The additive effects of density and harvest on adult survival of wolverines have relevance to the conservation and management of solitary carnivores with strong intrasexual territoriality, especially for species where combats among conspecifics can cause serious injury or even mortality.

The Norwegian red deer population (Cervus elaphus) was from the mid eighteenth to the early twentieth century drastically reduced in size and distribution but has the last century expanded both demographically and spatially. We have investigated genetic variation, differentiation and admixture in this spatially expanding ungulate population, using 14 microsatellites. The present genetic structure is moderate to strong with an average F ST = 0.08. Low M-ratios indicate loss of genetic variation in all localities and signals of a recent bottleneck was identified in 14 of 15 localities. Genetic distances between the localities indicate two main routes of dispersal during expansion, from the north–west and south–west, respectively. Bayesian assignment tests verify a break of the dataset in two, and demonstrate 99.9% probability for the existence of five sub-populations, which coincide well with five relict populations described by historic records. Computer simulations suggest that the observed genetic differentiation is recent rather than ancient, and that it may be explained by models of fragmentation or of founder events and subsequent merging rather than by models of recent bottlenecks in some particular demes within an ancient genetic structure.

Human population expansion and associated degradation of the habitat of many wildlife species cause loss of biodiversity and species extinctions. The small Simen Mountains National Park in Ethiopia is one of the last strongholds for the preservation of a number of afro-alpine mammals, plants and birds, and it is home to the rare endemic Walia ibex, Capra walie. The narrow distribution range of this species as well as potential competition for resources with livestock, especially with domestic goat, Capra hircus, may compromise its future survival. Based on a curated afro-alpine taxonomic reference library constructed for plant taxon identification, we investigated the diet of the Walia ibex and addressed the dietary overlap with domestic goat using DNA metabarcoding of faecal samples. Faeces of both species were collected from different localities in the National Park. We show that both species are browsers, with forbs, shrubs and trees comprising the largest proportion of their diet, supplemented by grasses. There was a considerable overlap in dietary preferences. Several of the preferred diet items of the Walia ibex (Alchemilla sp., Hypericum revolutum, Erica arborea and Rumex sp.) were also among the most preferred diet items of the domestic goat. These results indicate that there is potential for competition between the two species, especially during the dry season, when resources are limited. Our findings, in combination with the expected increase in domestic herbivores, suggest that management plans should consider the potential threat posed by domestic goats to ensure future survival of the endangered Walia ibex. published

Due to its conspicuous manifestations and its capacity to shape the configuration and dynamics of wild populations, territorial behavior has long intrigued ecologists. Territoriality and other animal interactions in situ have traditionally been studied via direct observations and telemetry. Here, we explore whether noninvasive genetic sampling, which is increasingly supplementing traditional field methods in ecological research, can reveal territorial behavior in an elusive carnivore, the wolverine (Gulo gulo). Using the locations of genotyped wolverine scat samples collected annually over a period of 12 years in central Norway, we test three predictions: (1) male home ranges constructed from noninvasive genetic sampling data are larger than those of females, (2) individuals avoid areas used by other conspecifics of the same sex (intrasexual territoriality), and (3) avoidance of same‐sex territories diminishes or disappears after the territory owner's death. Each of these predictions is substantiated by our results: sex‐specific differences in home range size and intrasexual territoriality in wolverine are patently reflected in the spatial and temporal configuration of noninvasively collected genetic samples. Our study confirms that wildlife monitoring programs can utilize the spatial information in noninvasive genetic sampling data to detect and quantify home ranges and social organization.

The southern Norwegian wolverine (Gulo gulo) population was considered functionally extinct in the 1960s but has partly recovered in recent years. Proper management of this population is highly dependent on reliable estimates of critical population parameters such as population size, sex ratio, immigration rate, and reproductive contribution from immigrants. We report on a large-scale population monitoring project assessing these parameters through genetic tagging of individuals, with feces as the source of DNA. Sixty-eight different individuals were detected among 147 successfully genotyped samples collected in 2000 and 2001. Sixty of these individuals were represented in the 2001 sample, which may be considered a minimum estimate of the population size. Almost 50% of these animals were sampled only once, however, indicating that the true population size may be markedly higher. Accordingly, a capture-recapture estimate based on the observed resampling rates suggested a population size of 89 wolverines (95% confidence interval (CI) = 74-104), which is approximately 35% higher than an estimate of 64 obtained from the number of active natal dens (95% CI = 46-95; p = 0.08). Indirect estimates of dispersal distances inferred from mother-offspring relationships sug- gested that wolverine males have the ability to disperse up to 500 km, a distance exceeding anything previously reported in the literature. Dispersal distances of more than 100 km were detected for females. Bayesian clus- tering analysis and subsequent assessment of individual relationships suggest that immigrants from northern Scandinavia have contributed and still contribute to the southern Norwegian gene pool, counteracting genetic erosion and reducing the risk of inbreeding depression. Additional sampling efforts will be undertaken during the coming years to allow for observations of population trends, immigration rate, and reproductive variance among individuals. Such data will provide an important basis for the design of an appropriate conservation plan for this small and vulnerable population.

The grey wolf ( Canis lupus ) was numerous on the Scandinavian peninsula in the early 19th century. However, as a result of intense persecution, the population declined dramatically and was virtually extinct from the peninsula by the 1960s. We examined historical patterns of genetic variability throughout the period of decline, from 1829 to 1979. Contemporary Finnish wolves, considered to be representative of a large eastern wolf population, were used for comparison. Mitochondrial DNA (mtDNA) variability among historical Scandinavian wolves was significantly lower than in Finland while Y chromosome variability was comparable between the two populations. This may suggest that long-distance migration from the east has been male-biased. Importantly though, as the historical population was significantly differentiated from contemporary Finnish wolves, the overall immigration rate to the Scandinavian peninsula appears to have been low. Levels of variability at autosomal microsatellite loci were high by the early 1800s but declined considerably towards the mid-20th century. At this time, approximately 40% of the allelic diversity and 30% of the heterozygosity had been lost. After 1940, however, there is evidence of several immigration events, coinciding with episodes of marked population increase in Russian Karelia and subsequent westwards migration.

The glacial-interglacial cycles of the upper Pleistocene have had a major impact on the recent evolutionary history of Arctic species. To assess the effects of these large-scale climatic fluctuations to a large, migratory Arctic mammal, we assessed the phylogeography of reindeer (Rangifer tarandus) as inferred from mitochondrial DNA (mtDNA) sequence variation in the control region. Phylogenetic relationships among haplotypes seem to reflect historical patterns of fragmentation and colonization rather than clear-cut relationships among extant populations and subspecies. Three major haplogroups were detected, presumably representing three separate populations during the last glacial. The most influential one has contributed to the gene pool of all extant subspecies and seems to represent a large and continuous glacial population extending from Beringia and far into Eurasia. A smaller, more localized refugium was most likely isolated in connection with ice expansion in western Eurasia. A third glacial refugium was presumably located south of the ice sheet in North America, possibly comprising several separate refugial populations. Significant demographic population expansion was detected for the two haplogroups representing the western Eurasian and Beringian glacial populations. The former apparently expanded when the ice cap retreated by the end of the last glacial. The large continuous one, in contrast, seems to have expanded by the end of the last interglacial, indicating that the warm interglacial climate accompanied by marine transgression and forest expansion significantly confined population size on the continental mainland. Our data demonstrate that the current subspecies designation does not reflect the mtDNA phylogeography of the species, which in turn may indicate that morphological differences among subspecies have evolved as adaptive responses to postglacial environmental change.

The fragmentation of populations is an increasingly important problem in the conservation of endangered species. Under these conditions, rare migration events may have important effects for the rescue of small and inbred populations. However, the relevance of such migration events to genetically depauperate natural populations is not supported by empirical data. We show here that the genetic diversity of the severely bottlenecked and geographically isolated Scandinavian population of grey wolves (Canis lupus), founded by only two individuals, was recovered by the arrival of a single immigrant. Before the arrival of this immigrant, for several generations the population comprised only a single breeding pack, necessarily involving matings between close relatives and resulting in a subsequent decline in individual heterozygosity. With the arrival of just a single immigrant, there is evidence of increased heterozygosity, significant outbreeding (inbreeding avoidance), a rapid spread of new alleles and exponential population growth. Our results imply that even rare interpopulation migration can lead to the rescue and recovery of isolated and endangered natural populations.

Carnivores are the top trophic level of food webs, eventually only superseded by man. Being at the top, they often require large spaces from which to obtain enough prey, a requirement that can create severemanagement complicationswhen they roamoutside protected areas or wildlife management zones. This can cause human-wildlife conflicts, usually resulting in the interface becoming a population sink for large predators (Woodroffe and Ginsberg, 1998). Human-carnivore relations are therefore often the proximate cause of the distribution and abundance of these predators today. But it is not only the carnivores’ relationship with man that limits their abundance. In spite of being at the top of the food web, carnivore populations can be regulated by other trophic levels, most commonly through food availability. Carnivores can also be regulated by intraguild competition, whereby the composition of a particular carnivore guild and their interspecific interactions determine the abundance of a given carnivore species within it. Furthermore, carnivores are potentially

Swayne's hartebeest (Alcelaphus buselaphus swaynei) is an endangered antelope that survives in four or five relict populations in Ethiopia. We examined the two main populations (Senkele and Nechisar) for mitochondrial (D-loop) and nuclear (microsatellite) variability in order to measure levels of genetic variation within the subspecies and degree of differentiation between populations. For comparison, we examined samples from a large population of red hartebeest (Alcelaphus buselaphus caama). Both swaynei and caama exhibited high levels of variation. There was significant differentiation between the populations of swaynei at Senkele and Nechisar, and gene diversity in Nechisar, the smaller of the two populations, was significantly lower than that in Senkele. Many mitochondrial haplotypes and microsatellite alleles present at high frequencies among the Senkele individuals were missing in Nechisar, suggesting that the translocation of animals from Senkele undertaken in 1974 did not contribute notably to the gene pool in Nechisar. Subsamples taken from Senkele in 1988 and 1995 showed a significant change in allele frequencies, a change that probably can be attributed to a massive population decline during this period. We recommend that both populations be protected in situ to maintain as much as possible of the diversity that exists within the taxon and that a breeding program be established. In spite of the earlier unsuccessful attempt, we argue that translocation of animals for enhancement of population size as well as genetic variation in Nechisar should be considered. Resumen: El alcelafo de Swayne (Alcelaphus buselaphus swaynei) es un antilope en peligro que sobrevive en cuatro o cinco poblaciones relicto en Etiopia. Examinamos la variabilidad mitocondrial (curva-D) y nuclear (microsatelite) en las dos principales poblaciones (Senkele y Nechisar) para medirlos niveles de variabilidad genetica en las subespecies y el nivel de diferenciacion entre poblaciones. Para comparacion, se examinaron muestras de una poblacion grande de alcelafo rojo (Alcelaphus buselaphus caama). Tanto swaynei como caama presentaron altos niveles de variacion. Hubo diferencia significativa entre las poblaciones de swaynei en Senkele y Nechisar, y la diversidad genica en Nechisar, la poblacion mas pequena, fue significativamente menor que la de Senkele. Muchos haplotipos mitocondriales y alelos microsatelitales presentes en frecuencias altas en Senkeles estuvieron ausentes en Nechisar, lo que sugiere que la translocacion de animales de Senkele en 1974 no contribuyo notablemente a la poza genica de Nechisar. Submuestras tomadas de Senkele en 1988 y 1995 mostraron un cambio significativo en la frecuencia de alelos, cambio que se puede atribuir a la declinacion poblacional masiva durante ese periodo. Recomendamos que las poblaciones se protejan in situ para mantener la mayor diversidad posible dentro del taxon y que se establezca un programa de reproduccion. A pesar del infructuoso intento anterior, se debe considerar la translocacion de animales para incrementar el tamano poblacional y la variabilidad genetica en Nechisar.

1 Animal Genetic Resource Department, Institute of Biodiversity Conservation, Addis Ababa, Ethiopia 2 Laboratoire d’Ecologie Alpine, CNRS UMR 5553, Universite Joseph Fourier, Grenoble, France 3 Department of Environmental Sciences, University of Milano-Bicocca, Milano, Italy 4 Natural Resources Faculty, University of Guilan, Guilan, Iran 5 Environmental Sciences Department, Gorgan University of Agriculture and Natural Resources, Gorgan, Iran 6 Centro de Investigacao em Biodiversidade e Recursos Geneticos (CIBIO-UP), Universidade do Porto, Vairao, Portugal 7 Division of Biological Sciences, University of Montana, Missoula, MT, USA 8 Norwegian Institute for Nature Research, Trondheim, Norway

The globally endangered snow leopard Panthera uncia is sparsely distributed throughout the rugged mountains in Asia. Its habit of preying on livestock poses a main challenge to management. In the remote Phu valley in northern Nepal, we obtained reliable information on livestock losses and estimated predator abundance and diet composition from DNA analysis and prey remains in scats. The annual diet consisted of 42%livestock. Among the wild prey, bharal (blue sheep/ naur) Pseudois nayaur was by far the most common species (92%). Two independent abundance estimates suggested that there were six snow leopards in the valley during the course of our study. On average, each snow leopard killed about one livestock individual and two bharal permonth. Predation loss of livestock estimated from prey remains in scats was 3.9%, which was in concordance with village records (4.0%). From a total count of bharal, the only large natural prey in the area and occurring at a density of 8.4 animals/km2 or about half the density of livestock, snow leopards were estimated to harvest 15.1% of the population annually. This predation rate approaches the natural, inherent recruitment rate of this species; in Phu the proportion of kids was estimated at 18.4%. High livestock losses have created a hostile attitude against the snow leopard and mitigation measures are needed. Among innovative management schemes now being implemented throughout the species’ range, compensation and insurance programmes coupled with other incentive measures are encouraged, rather than measures to reduce the snow leopard’s access to livestock. In areas like the Phu valley, where the natural prey base consists mainly of one ungulate species that is already heavily preyed upon, the latter approach, if implemented, will lead to increased predation on this prey, which over time may suppress numbers of both prey and predator. Snow leopard Panthera uncia predation on livestock and wild prey in a mountain valley in northern Nepal: implications for conservation management

Many East African mountains are characterized by an exceptionally high biodiversity. Here we assess the hypothesis that climatic fluctuations during the Plio-Pleistocene led to ecological fragmentation with subsequent genetic isolation and speciation in forest habitats in East Africa. Hypotheses on speciation in savannah lineages are also investigated. To do this, mitochondrial DNA sequences from a group of bush crickets consisting of both forest and savannah inhabiting taxa were analysed in relation to Plio-Pleistocene range fragmentations indicated by palaeoclimatic studies. Coalescent modelling and mismatch distributions were used to distinguish between alternative biogeographical scenarios. The results indicate two radiations: the earliest one overlaps in time with the global spread of C4 grasslands and only grassland inhabiting lineages originated in this radiation. Climatically induced retraction of forest to higher altitudes about 0.8 million years ago, promoting vicariant speciation in species inhabiting the montane zone, can explain the second radiation. Although much of the biodiversity in East Africa is presently threatened by climate change, past climatic fluctuations appear to have contributed to the species richness observed in the East African hot spots. Perceiving forests as centres of speciation reinforces the importance of conserving the remaining forest patches in the region.

International audience; Knowledge about the phylogenetic history, genetic variation and ecological requirements of a species is important for its conservation and management. Unfortunately, for many species this information is lacking. Here we use multiple approaches (phylogenetics, population genetics and ecological modelling) to evaluate the evolutionary history and conservation status of Capra walie, an endangered flagship species of wild goat endemic to Ethiopia. The analysis of mitochondrial cytochrome b and Y-chromosome DNA sequences suggests that C. walie forms a monophyletic clade with Capra nubiana, but potentially has been isolated for up to 0.8 million years from this closely related species. Microsatellite DNA analyses show that C. walie has very low genetic variation (mean heterozygosity=0.35) compared with other endangered mammals. This reduced variation likely derives from a prolonged demographic decline and small effective population size. Ecological niche modelling using the bioclimatic features of habitats occupied by C. walie, suggests ecological differences between C. walie and C. nubiana, and identifies the areas most suitable for future reintroductions of C. walie. The genetic and bioclimatic data suggest that C. walie is distinct and requires immediate conservation actions including genetic monitoring and reintroductions to establish independent populations. This study illustrates how combining noninvasive sampling along with genetic and ecological (bioclimatic) approaches can help assess conservation status of poorly known species.

1. Wolves Canis lupus L. recolonized the boreal forests in the southern part of the Scandinavian peninsula during the late 1990s, but so far there has been little attention to its effect on ecosystem functioning. Wolf predation increases the availability of carcasses of large prey, especially moose Alces alces L., which may lead in turn to a diet switch in facultative scavengers such as the wolverine Gulo gulo L. 2. Using 459 wolverine scats collected during winter-spring 2001-04 for DNA identity and dietary contents, we compared diet inside and outside wolf territories while controlling for potential confounding factors, such as prey density. We tested the hypothesis that wolverine diet shifted towards moose in the presence of wolves, while taking into account possible sexual segregation between the sexes. Occurrence of reindeer, moose and small prey was modelled against explanatory covariates using logistic mixed-effects models. Furthermore, we compared diet composition and breadth among habitats and sexes. 3. Occurrence of reindeer, moose and small prey in the diet varied with prey availability and habitat. As expected, diet contained more moose and less reindeer and small prey in the presence of wolves. Their diet in tundra consisted of 40% reindeer Rangifer tarandus L., 39% moose and 9% rodents. In forest with wolf, their diet shifted to 76% moose, 18% reindeer and 5% rodents; compared to 42% moose, 32% reindeer and 15% rodents in forest without wolf. This diet switch could not be explained by higher moose density in wolf territories. Female diet consisted of more small prey than for males, but there was a tendency for females to use the highly available moose carrion opportunistically and to hunt less on small prey within wolf territories. 4. Our study highlights how wolves increase scavenging opportunities for wolverines, and how sexual differences in diet may also apply to large scavengers. Due to their more restricted home range, female wolverines are forced to rely more on hunting small prey. The relatively high occurrence of wolf kills, however, forms an important food source to wolverines in this area. The recolonization of wolves may therefore have contributed to the consequent recolonization of wolverines into the same area.

Although there is little doubt that the domestication of mammals was instrumental for the modernization of human societies, even basic features of the path towards domestication remain largely unresolved for many species. Reindeer are considered to be in the early phase of domestication with wild and domestic herds still coexisting widely across Eurasia. This provides a unique model system for understanding how the early domestication process may have taken place. We analysed mitochondrial sequences and nuclear microsatellites in domestic and wild herds throughout Eurasia to address the origin of reindeer herding and domestication history. Our data demonstrate independent origins of domestic reindeer in Russia and Fennoscandia. This implies that the Saami people of Fennoscandia domesticated their own reindeer independently of the indigenous cultures in western Russia. We also found that augmentation of local reindeer herds by crossing with wild animals has been common. However, some wild reindeer populations have not contributed to the domestic gene pool, suggesting variation in domestication potential among populations. These differences may explain why geographically isolated indigenous groups have been able to make the technological shift from mobile hunting to large-scale reindeer pastoralism independently.

The identification of hybrids is often a subject of primary concern for the development of conservation and management strategies, but can be difficult when the hybridizing species are closely related and do not possess diagnostic genetic markers. However, the combined use of mitochondrial DNA (mtDNA), autosomal and Y chromosome genetic markers may allow the identification of hybrids and of the direction of hybridization. We used these three types of markers to genetically characterize one possible wolf-dog hybrid in the endangered Scandinavian wolf population. We first characterized the variability of mtDNA and Y chromosome markers in Scandinavian wolves as well as in neighboring wolf populations and in dogs. While the mtDNA data suggested that the target sample could correspond to a wolf, its Y chromosome type had not been observed before in Scandinavian wolves. We compared the genotype of the target sample at 18 autosomal microsatellite markers with those expected in pure specimens and in hybrids using assignment tests. The combined results led to the conclusion that the animal was a hybrid between a Scandinavian female wolf and a male dog. This finding confirms that inter-specific hybridization between wolves and dogs can occur in natural wolf populations. A possible correlation between hybridization and wolf population density and disturbance deserves further research.