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For many species, the ability to rapidly adapt to changes in seasonality is essential for long-term survival. In the Arctic, seasonal moulting is a key life-history event that provides year-round camouflage and thermal protection. However, increased climatic variability of seasonal events can lead to phenological mismatch. In this study, we investigated whether winter-white (white morph) and winter-brown (blue morph) Arctic foxes (Vulpes lagopus) could adjust their winter-to-summer moult to match local environmental conditions. We used camera trap images spanning an eight-year period to quantify the timing and rate of fur change in a polymorphic subpopulation in south-central Norway. Seasonal snow cover duration and temperature governed the phenology of the spring moult. We observed a later onset and longer moulting duration with decreasing temperature and longer snow season. Additionally, white foxes moulted earlier than blue in years with shorter periods of snow cover and warmer temperatures. These results suggest that phenotypic plasticity allows Arctic foxes to modulate the timing and rate of their spring moult as snow conditions and temperatures fluctuate. With the Arctic warming at an unprecedented rate, understanding the capacity of polar species to physiologically adapt to a changing environment is urgently needed in order to develop adaptive conservation efforts. Moreover, we provide the first evidence for variations in the moulting phenology of blue and white Arctic foxes. Our study underlines the different intraspecific selective pressures that can exist in populations where several morphs co-occur, and illustrates the importance of integrating morph-based differences in future management strategies of such polymorphic species. camera trapping, camouflage, climate change, colour polymorphism, phenological mismatch, phenotypic plasticity

Accelerating climate change is causing severe habitat fragmentation in the Arctic, threatening the persistence of many cold-adapted species. The Scandinavian arctic fox (

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.

Genetic drift can dramatically change allele frequencies in small populations and lead to reduced levels of genetic diversity, including loss of segregating variants. However, there is a shortage of quantitative studies of how genetic diversity changes over time in natural populations, especially on genome-wide scales. Here, we analyzed whole-genome sequences from 76 wolves of a highly inbred Scandinavian population, founded by only one female and two males, sampled over a period of 30 yr. We obtained chromosome-level haplotypes of all three founders and found that 10%-24% of their diploid genomes had become lost after about 20 yr of inbreeding (which approximately corresponds to five generations). Lost haplotypes spanned large genomic regions, as expected from the amount of recombination during this limited time period. Altogether, 160,000 SNP alleles became lost from the population, which may include adaptive variants as well as wild-type alleles masking recessively deleterious alleles. Although not sampled, we could indirectly infer that the two male founders had megabase-sized runs of homozygosity and that all three founders showed significant haplotype sharing, meaning that there were on average only 4.2 unique haplotypes in the six copies of each autosome that the founders brought into the population. This violates the assumption of unrelated founder haplotypes often made in conservation and management of endangered species. Our study provides a novel view of how whole-genome resequencing of temporally stratified samples can be used to visualize and directly quantify the consequences of genetic drift in a small inbred population.

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.

Spatial capture-recapture modelling (SCR) is a powerful tool for estimating densities, population size and space use of elusive animals. Here, we applied SCR modeling 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 northern study area (mean = 0.04 foxes per km2 [95%CI: 0.02-0.09] in 2016, 0.09 [0.05-0.18] in 2017 and 0.07 [0.04-0.13] in 2018) compared to the southern study area (0.16 [0.09-0.26] in 2017 and 0.10 [0.07-0.16] in 2018). We found a positive effect of forest cover on density in the northern, 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, but could also be due to climatic differences (e.g., winter severity) between the two areas. Estimated mean home range size in the northern study area was 45 km2 [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 modeling 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.

Age at first reproduction constitutes a key life-history trait in animals and is evolutionarily shaped by fitness benefits and costs of delayed versus early reproduction. The understanding of how intrinsic and extrinsic changes affects age at first reproduction is crucial for conservation and management of threatened species because of its demographic effects on population growth and generation time. For a period of 40 years in the Scandinavian wolf (Canis lupus) population, including the recolonization phase, we estimated age at first successful reproduction (pup survival to at least three weeks of age) and examined how the variation among individuals was explained by sex, population size (from 1 to 74 packs), primiparous or multiparous origin, reproductive experience of the partner and inbreeding. Median age at first reproduction was 3 years for females (n = 60) and 2 years for males (n = 74), and ranged between 1 and 8–10 years of age (n = 297). Female age at first reproduction decreased with increasing population size, and increased with higher levels of inbreeding. The probability for males to reproduce later first decreased, reaching its minimum when the number of territories approached 40–60, and then increased with increasing population size. Inbreeding for males and reproductive experience of parents and partners for both sexes had overall weak effects on age at first reproduction. These results allow for more accurate parameter estimates when modelling population dynamics for management and conservation of small and vulnerable wolf populations, and show how humans through legal harvest and illegal hunting influence an important life-history trait like age at first reproduction.

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.

In wide ranging species, portions of the same population often fall within different administrative jurisdictions; where different regulations apply. The same species can be fully protected or heavily harvested on different sides of a border. This can generate a source–sink dynamic from the areas with lower to those with higher mortality, a process known as compensatory immigration. We tested this hypothesis on the wolverine (Gulo gulo) population of southern Scandinavia, which is shared between two countries: Sweden and Norway. Wolverines are fully protected in Sweden, but subject to intensive population regulation in Norway. Using non-invasive genetic sampling and capture–recapture modeling, we analyzed the dynamics of wolverine survival and emigration patterns between 2002 and 2013. Wolverines in Norway experienced a lower survival than in Sweden. Migration across the national border was directed towards movements from Sweden to Norway. There was a functional relationship between harvest rate in Norway and emigration rates across the national border, both at the individual and population level, thus confirming the compensatory immigration hypothesis. Contrasting management regimes within the same population can generate undesired demographic and spatial dynamics, jeopardize conservation goals on the two sides of a border, and reduce the efficiency of management actions. This calls for the adoption of a coordinate population approach in large carnivore conservation and management. Failing to do so can cause a waste of the already limited resources allocated for large carnivore conservation, and it might hinder effective conflict mitigation.

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.

Genetic approaches have proved valuable to the study and conservation of endangered populations, especially for monitoring programs, and there is potential for further developments in this direction by extending analyses to the genomic level. We assembled the genome of the wolverine (Gulo gulo), a mustelid that in Scandinavia has recently recovered from a significant population decline, and obtained a 2.42 Gb draft sequence representing85% of the genome and including21,000 protein-coding genes. We then performed whole-genome resequencing of 10 Scandinavian wolverines for population genomic and demographic analyses. Genetic diversity was among the lowest detected in a red-listed population (mean genome-wide nucleotide diversity of 0.05%). Results of the demographic analyses indicated a long-term decline of the effective population size (N

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.

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.

The Terai is one of the world's most spectacular landscapes, encompassing parts of Nepal and northern India. This area used to harbour large and continuous populations of charismatic species like elephants, tigers and rhinoceros. However, recent habitat fragmentation reduced these populations into small, partially or completely isolated remnants. The largest of these fragments in Nepal is the Bardia National Park. Here, the elephant population was functionally extinct in the early 1970s and -80s, but was rescued by a considerable number of immigrants in 1994. In order to assess population size, sex ratio, age structure, and levels of genetic variation, we carried out non-invasive genetic sampling, using elephant dung as the source of DNA. A capture-mark-recapture estimate of population size suggested that there were 57 individuals in the study area, which agrees well with field observations. Notably, a strongly male-biased sex ratio was evident among sub-adult individuals. This observation suggests the presence of sub-adult immigrants in the population, which was supported by formal migrant detection analysis. Genetic variation was quite high and the evidence for male immigrants suggests that there are good prospects for maintenance of genetic diversity. A decade ago a large-scale project was initiated in the Terai region to link remaining populations of large mammals through dispersal corridors. The program is basically founded on the assumption that habitat fragments are isolated with little or no migration between them. Our results indicate that this may not be the case, at least not for the Asian elephant in western Nepal, which therefore reduces the alleged extinction risk from genetic erosion and stochastic demographic events.

The timing and origin of reindeer domestication has been highly debated. Recent molecular analyses show several mitochondrial lineages of domestic reindeer across Eurasia suggesting different origins of Fennoscandian and Siberian domestic reindeer. In order to investigate the origin of domestic Fennoscandian reindeer, we sequenced a mitochondrial control region fragment of 68 ancient reindeer remains from archaeological sites in Finnmark, the major county for extant reindeer husbandry in Norway, spanning from ca. BC 3400 to AD 1800. The majority of the Stone and Iron Age reindeer assemblages in Finnmark are from settlements in the eastern part of the county, in the Varangerfjord area. The reindeer remains from these settlements show affiliation to the large and complex Beringian haplotype cluster, found in extant reindeer from the Kola Peninsula to north-eastern Russia. A distinct haplotype shift is observed in the late medieval period, when the typical haplotype signatures of extant domestic Fennoscandian reindeer appeared in coastal regions of both eastern and western Finnmark. These haplotypes were not found among the Stone and Iron Age wild reindeer samples of Finnmark, suggesting that this population was not ancestral to extant domestic reindeer of Fennoscandia.

Despite the importance of the reindeer husbandry in the subsistence of many northern cultures the origin and spread of domestic reindeer is still highly debated. Recent analyses of mitochondrial DNA in reindeer herds across Eurasia revealed distinct geographic structures of domestic reindeer in Fennoscandia and Russia, pointing towards independent origins of domestic reindeer in these areas. A high degree of haplotype sharing between the extant wild reindeer population in the Hardangervidda mountain region in southern Norway and the domestic herds in Fennoscandia indicate that this population could have contributed in the early domestication or augmented the domestic population. However, genetic analyses of excavated reindeer remains from the early medieval period from the Hardangervidda region revealed no haplotype sharing with extant domestic reindeer, demonstrating that reindeer from Hardangervidda did not contribute in the early domestication process in Fennoscandia. The substantial temporal genetic alteration observed in this population is related to introgression of domestic reindeer into the wild gene pool in the 19th century during periods when reindeer husbandry was practiced in this mountain region.

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.

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

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.

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.