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Climate change increases the risk of severe alterations to essential wildlife habitats. The Arctic fox (Vulpes lagopus (Linnaeus, 1758)) uses dens as shelters against cold temperatures and predators. These dens, needed for successful reproduction, are generally dug into the active layer on top of permafrost and reused across multiple generations. We assessed the vulnerability of Arctic fox dens to the increasing frequency of geohazards (thaw settlement, mass movements, and thermal erosion) that is arising from climate change. On Bylot Island (Nunavut, Canada) we developed, and calculated from field observations, a qualitative vulnerability index to geohazards for Arctic fox dens. Of the 106 dens studied, 14% were classified as highly vulnerable, whereas 17% and 69% had a moderate and low vulnerability, respectively. Vulnerability was not related to the probability of use for reproduction. Although climate change will likely impact Arctic fox reproductive dens, such impact is not a major threat to foxes of Bylot Island. Our research provides the first insights into the climate-related geohazards potentially affecting Arctic fox ecology in the next decades. The developed method is flexible and could be applied to other locations or other species that complete their life cycle in permafrost regions.

Climate-caused changes in prey abundance may alter predator–prey dynamics in the Arctic food web. Lemmings (Dicrostonyx spp.) are important prey for Arctic foxes (Vulpes lagopus) and their annual population fluctuations drive fox reproduction, creating strongly linked predator–prey population cycles. Winter diet directly impacts Arctic fox reproductive success, but winter prey diversity on the tundra is low. Strategies such as using the marine environment to scavenge seals have allowed Arctic foxes to persist during years of low lemming abundance. However, warming winters have decreased snowpack quality, preventing lemmings from reaching their previous high abundances, which may reduce their impact on predator dynamics. We investigated Arctic fox dietary response to lemming abundance by reconstructing Arctic fox winter diet in the low Arctic. Next-generation sequencing of fecal DNA, from samples (n = 627) collected at dens in winters of 2011–2018, identified prey both from terrestrial and marine environments. Despite lemming cycle damping, Arctic foxes still increased lemming consumption during years of higher lemming abundance. Alternative prey such as marine resources were consumed more during years of low lemming abundance, with up to 45% of samples containing marine resources in low lemming years. In addition, Arctic foxes consumed high proportions of meadow voles (Microtus pennsylvanicus), which may represent a new alternative prey, suggesting climate change may be creating new foraging opportunities. Changes in prey abundance illustrate how climate-caused disturbances are altering Arctic food-web dynamics. Dietary flexibility and availability of alternative prey may become increasingly important for Arctic predators as the Arctic continues to change.

Arctic fox (Vulpes lagopus) and its tundra habitat are a unique system for the study of rabies virus epidemics. Contrary to theoretical calculations reporting a critical density (KT) of approximate...

As the interest for nature-based tourism activities increases, it is important to provide evidence-based guidelines for wildlife-human interactions to minimize the disturbance caused to wildlife. I ...

Supplementary feeding is often used as a conservation tool to reverse the decline of food‐limited populations. The arctic fox (Vulpes lagopus) is one of the most endangered mammals in Norway and has been the target of several conservation initiatives for almost 3 decades, including supplementary feeding. To measure and improve the efficiency of supplementary feeding as a conservation action, we used passive integrated transponder (PIT)‐tags in arctic foxes and 6 feeding stations equipped with PIT‐tag readers to monitor individual use of supplemental food between 2013 and 2018. We tested hypotheses about the potential influence of temporal and spatial patterns, individual characteristics (i.e., age, sex, reproductive status), and food abundance (abundance of small rodents and amount of food filled) on the frequency and intensity of use of supplementary feeding stations by arctic foxes. The feeding stations were visited ≥1 time by 196 PIT‐tagged individuals. We detected 54% of juveniles born in the study area between 2013 and 2017 at the feeding stations. More arctic foxes used the feeding stations during the pre‐breeding period than during the other seasons, and the visits occurred mostly at night. The closest feeding station to each natal den was systematically used by the established pair and by the juveniles born at this den. Juveniles did not use the feeding stations more than adult foxes. Older foxes, and breeding adults, visited the feeding stations more than younger and non‐breeding adults. Foxes used feeding stations more intensively when prey was scarce and with greater amounts of supplemental food. This study highlights that supplemental feeding is important for breeding adults, especially in periods of low prey abundance. Understanding the use of feeding stations will contribute to the optimization of supplemental feeding as a conservation action and help wildlife managers to carefully plan and manage its discontinuation.

Isolation at small population size can reduce individual fitness and impede population growth caused by inbreeding and genetic drift (i.e. inbreeding depression). Inbreeding depression can however be circumvented by gene flow from unrelated individuals through masking of recessive deleterious alleles and contribute to population persistence (i.e. genetic rescue). Studying these processes in natural populations across generations and under fluctuating environmental conditions however comes with major challenges. Several gaps in the knowledge thus remain regarding causes and consequences of inbreeding depression and genetic rescue in the wild. Using long term data on life history traits, combined with traditional population genetics and novel genomic techniques, we explored the dynamics of inbreeding and gene flow in the highly fragmented arctic fox (Vulpes lagopus) population in Sweden. This thesis mainly focused on the southernmost subpopulation (Helagsfjällen), previously documented to suffer from inbreeding depression. Construction of a genetically verified pedigree (Chapter I and II) revealed that gene flow from three outbred male foxes released from a captive breeding station in Norway resulted in genetic rescue, expressed as elevated first year survival and breeding success in immigrant first generation offspring (F1; Chapter I). However, the rescue effect likely only lasted for one single generation, as we found no selective advantage in later descendants of immigrants (Chapter II and IV). Whole genome sequencing of a subset of individuals from the same subpopulation showed that some immigrant F2 and F3 individuals were highly inbred despite the recent outbreeding events (Chapter III). Identification of putative deleterious variation within coding regions suggested that the immigrants introduced a large number of strongly deleterious alleles which were absent from the native gene pool (Chapter IV and V). Expression of the deleterious variation introduced may explain the low persistence of genetic rescue. We also found a negative relationship between the amount of homozygous strongly deleterious mutations and individual fitness (Chapter IV) and may be an important cause of inbreeding depression in the Swedish arctic fox. Finally, when replicating the study of genomic consequences of inbreeding and gene flow, by including an additional Swedish subpopulation (Vindelfjällen) located further north, we found contrasting patterns between the two subpopulations. While inbreeding decreased in both Helagsfjällen and Vindelfjällen following immigration, the proportion of deleterious variation increased in Helagsfjällen but not in Vindelfjällen. A potential explanation could be more regular gene flow between northern located subpopulations compared to the more geographically isolated population in Helagsfjällen, which may instead have purged a subset of strongly deleterious variation pre immigration. The results from this thesis highlight the transient nature of genetic rescue and the importance to study fitness and genetic effects of gene flow across several generations, as immigration could have negative consequences that are not manifested initially. Finally, as the effects of gene flow can be highly context dependent, demographic histories and functional genetic variation in both source and target populations should be considered before making translocation decisions for conservation purposes.

Translocations are a widespread approach to conserve threatened populations. Given the rapid decline and genetic deterioration of many natural populations, translocations are expected to become even more common in the future. The success of translocations is, however, dependent on multiple context-dependent factors, such as demographic and genetic status, habitat quality and animal behaviour. The Scandinavian arctic fox (Vulpes lagopus) exists in a small, fragmented population that is demographically vulnerable and exposed to inbreeding depression. In the early 2000 s, releases of arctic foxes from the Norwegian Captive Breeding Programme were initiated with the purpose of reintroducing populations to formerly inhabited areas and promoting connectivity. Since 2008/2009, 61 foxes have been released in Junkeren, Norway to re-establish an unoccupied area. We used a combination of field observations and microsatellite genotyping from the release site and two neighbouring subpopulations to investigate (i) the probability of establishment and reproduction for released foxes at the release site and in neighbouring subpopulations, and (ii) the impact on litter size and genetic composition in the recipient populations. Results showed that 18% of the released foxes were established at the release site, or in neighbouring subpopulations and 11.5% reproduced successfully. The extent of post-release dispersal into neighbouring subpopulations was also relatively high (11.5%). During the study period, the number of litters more than doubled in the subpopulations with released foxes contributing 29.5% to this increase, but no clear effect of immigration on litter size was found. There was a slight increase in genetic variation in one of the subpopulations, and a significant decline in genetic divergence between subpopulations. We conclude that despite extensive releases, demographic and genetic effects were highly context-dependent. This study highlights the challenges of reinforcement programmes in small populations and reintroductions to unoccupied sites, especially for highly mobile species in a fragmented landscape. conservation; immigration; translocation; genetic variation; demography; arctic fox; Vulpes lagopus; captive breeding; conservation breeding.

This is a pre-print of an article published in Oecologia. The final authenticated version is available online at: https://doi.org/10.1007/s00442-019-04580-0 Identifying resources driving long-term trends in predators is important to understand ecosystem changes and to manage populations in the context of conservation or control. The arctic fox population in Iceland has increased steadily over a period of 30 years, an increase that has been attributed to an overall increase in food abundance. We hypothesized that increasing populations of geese or seabirds were driving this growth. We analyzed stable isotopes in a long-term series of collagen samples to determine the role of these different resources. The isotopic signatures of arctic foxes differed consistently between coastal and inland habitats. While δ15N displayed a non-linear change over time with a slight increase in the first part of the period followed by a decline in both habitats, δ13C was stable. Stable isotope mixing models suggested that marine resources and rock ptarmigan were the most important dietary sources, with marine resources dominating in coastal habitats and rock ptarmigan being more important inland. Our results suggest that seabirds may have been driving the arctic fox population increase. The rapidly increasing populations of breeding geese seem to have played a minor role in arctic fox population growth, as rock ptarmigan was the most important terrestrial resource despite a considerable decrease in their abundance during recent decades. This study shows that a long-term population trend in a generalist predator may have occurred without a pronounced change in main dietary resources, despite ongoing structural changes in the food web, where one species of herbivorous birds increased and another decreased.

Predation shapes communities through consumptive and non‐consumptive effects. In the latter case, prey respond to perceived predation risk through proactive or reactive risk management strategies occurring at different spatial and temporal scales. The predator–prey space race and landscape of fear concepts are useful to better understand how predation risk affects prey behavioral decisions and distribution. We assessed predation risk effects in a terrestrial Arctic community, where the arctic fox is the main predator of ground‐nesting birds. Using high‐frequency GPS data, we estimated a predator activity landscape corresponding to fox space use patterns and validated with an artificial prey experiment that this predator activity landscape correlated with the predation risk landscape. We then investigated the effects of the fox activity landscape on multiple prey species, by assessing the anti‐predator behavior of a main prey (snow goose) actively searched for by foxes, and the nest distribution of several incidental prey species. We first found that snow geese showed a stronger level of nest defense in areas highly used by foxes, possibly responding with a reactive strategy to variation in predation risk. Then, nests of incidental prey reproducing in habitats easily accessed by foxes had a lower probability of occurrence in areas highly used by foxes, suggesting these birds may use a proactive risk management strategy by shifting their distribution away from risky areas. For incidental prey species nesting in microhabitat refuges difficult to access by foxes, probability of nest occurrence was independent of predation risk in the surrounding area, as they avoid risk at a finer spatial scale. By tracking all individuals of the dominant predator species in our study area, we demonstrated the value of using predator space use patterns to infer spatial variation in predation risk. Overall, we highlight the diversity of risk management strategies in prey sharing a common predator, hence refining our understanding of the mechanisms driving species distribution and community structure.

Background Biologging now allows detailed recording of animal movement, thus informing behavioural ecology in ways unthinkable just a few years ago. In particular, combining GPS and accelerometry allows spatially explicit tracking of various behaviours, including predation events in large terrestrial mammalian predators. Specifically, identification of location clusters resulting from prey handling allows efficient location of killing events. For small predators with short prey handling times, however, identifying predation events through technology remains unresolved. We propose that a promising avenue emerges when specific foraging behaviours generate diagnostic acceleration patterns. One such example is the caching behaviour of the arctic fox (Vulpes lagopus), an active hunting predator strongly relying on food storage when living in proximity to bird colonies. Methods We equipped 16 Arctic foxes from Bylot Island (Nunavut, Canada) with GPS and accelerometers, yielding 23 fox-summers of movement data. Accelerometers recorded tri-axial acceleration at 50 Hz while we obtained a sample of simultaneous video recordings of fox behaviour. Multiple supervised machine learning algorithms were tested to classify accelerometry data into 4 behaviours: motionless, running, walking and digging, the latter being associated with food caching. Finally, we assessed the spatio-temporal concordance of fox digging and greater snow goose (Anser caerulescens antlanticus) nesting, to test the ecological relevance of our behavioural classification in a well-known study system dominated by top-down trophic interactions. Results The random forest model yielded the best behavioural classification, with accuracies for each behaviour over 96%. Overall, arctic foxes spent 49% of the time motionless, 34% running, 9% walking, and 8% digging. The probability of digging increased with goose nest density and this result held during both goose egg incubation and brooding periods. Conclusions Accelerometry combined with GPS allowed us to track across space and time a critical foraging behaviour from a small active hunting predator, informing on spatio-temporal distribution of predation risk in an Arctic vertebrate community. Our study opens new possibilities for assessing the foraging behaviour of terrestrial predators, a key step to disentangle the subtle mechanisms structuring many predator–prey interactions and trophic networks.

Despite the availability of ever more data on population trends in Arctic waders, we still have limited insight into the factorsinfluencing reproductive success and its resulting contribution to population trajectories. Here, we present data from Zackenbergin high Arctic Greenland that show no changes in timing of egg laying nor clutch size over more than two decades.With variation among species, we found earlier egg laying in seasons with earlier snowmelt, more invertebrates during thepre-laying period, earlier appearance of invertebrates and higher June temperatures. In addition, we found larger clutch sizeswith earlier snowmelt, and lower clutch sizes with later nest initiation dates, together with increasing variability in clutchsize during the study years in one species. Nest success was lower in late breeding seasons but seemed to increase withseason progress, while nest success was much lower during the lasts 18 years of the study period than during the first part ofthe period. The reason for this could be both increased Arctic fox activity on the tundra and increased researcher activity atnests, which leaves olfactory clues for the foxes. Numbers of juvenile Dunlins produced seem to be negatively correlated withour index of fox activity on the tundra during nesting and fledging, whereas we found no correlations between fox activityand lemming abundance. The effects of severe events in a number of years are described, including a next to non-breedingyear in 2018. Our study adds to our understanding that the Arctic is not a homogeneous biome but may show quite differentregional patterns of climate change effects.

Climate change has different and sometimes divergent effects on terrestrial and marine food webs, and in coastal ecosystems, these effects are tightly interlinked. Responses of opportunistic coastal predators and scavengers to climate change may thus be complex and potentially highly flexible, and can simultaneously serve as indicators of, and have profound impacts on, lower trophic levels. Gaining mechanistic understanding of these responses is therefore important, but often not feasible due to lack of longterm data from marked individuals. Here, we used a Bayesian integrated population model (IPM) to elucidate the effects of arctic warming and concurrent changes in terrestrial and marine resource availability on population dynamics of the opportunistic arctic fox (Vulpes lagopus) in Svalbard. Joint analysis of four types of data (den survey, age-at-harvest, placental scars, mark-recovery) revealed relatively stable population size and age structure over the last 22 yr (1997–2019) despite rapid environmental change linked to climate warming. This was related to the fact that terrestrial resources (reindeer carcasses, geese) became more abundant while the availability of marine resources (seal pups/carrion) decreased, and was driven by divergent trends in different vital rates (e.g., increased pregnancy rate but decreased pup survival). Balanced contributions of survival vs. reproduction and of immigration vs. local demography further stabilized population size. Our study thus sheds light on the mechanisms underlying population dynamics of opportunistic carnivores exploiting terrestrial and marine resources and suggests that exploitation of resources across different ecosystems can buffer predators against climate change. Additionally, it highlights the large potential of IPMs as tools to understand and predict the effects of environmental change on wildlife populations, even when data on marked individuals are sparse. arctic fox; cause-specific mortality; demography; harvest; immigration; integrated population model; marine resources; population dynamics; sea ice; terrestrial resources; transient life table response experiment Open Access CC-BY

Climate change can impact ecosystems by reshaping the dynamics of resource exploitation for predators and their prey. Alterations of these pathways could be especially intense in ecosystems characterized by a simple trophic structure and rapid warming trends, such as in the Arctic. However, quantifying the multiple direct and indirect pathways through which climate change is likely to alter trophic interactions and their relative strength remains a challenge. Here, we aim to identify direct and indirect causal mechanisms driven by climate affecting predator-prey interactions of species sharing a tundra food web. We based our study on relationships between one Arctic predator (Arctic fox) and its two main prey - lemmings (preferred prey) and snow geese (alternate prey) - which are exposed to variable local and regional climatic factors across years. We used a combination of models mapping multiple causal links among key variables derived from a long-term dataset (21 years). We obtained several possible scenarios linking regional climate factors (Arctic oscillations) and local temperature and precipitation to the breeding of species. Our results suggest that both regional and local climate factors have direct and indirect impacts on the breeding of foxes and geese. Local climate showed a positive causal link with goose nesting success, while both regional and local climate displayed contrasted effects on the proportion of fox breeding. We found no impact of climate on lemming abundance. We observed positive relationships between lemming, fox and goose reproduction highlighting numerical and functional responses of fox to the variability of lemming abundance. Our study measures causal links and strength of interactions in a food web, quantifying both numerical response of a predator and apparent interactions between its two main prey. These results improve our understanding of the complex effects of climate on predator-prey interactions and our capacity to anticipate food web response to ongoing climate change.Les changements climatiques peuvent avoir un impact sur les écosystèmes au travers des modifications de la dynamique d'exploitation des ressources par les prédateurs et leurs proies. Dans le cas de l'Arctique, caractérisée par un réseau trophique simple et une sensibilité marquée au réchauffement climatique, l'altération de ces relations trophiques pourrait être particulièrement importante. Cependant, la quantification des nombreux liens directs et indirects à travers lesquels les changements climatiques peuvent affecter les interactions trophiques demeure un défi. Notre objectif est d'identifier les mécanismes causaux directs et indirects, sous-tendus par le climat, affectant les interactions prédateur-proie au sein d'un réseau trophique au cœur de la toundra. Notre étude se base sur les relations entre un prédateur (renard arctique) et ses deux proies principales -le lemming (proie préférée) et la grande oie des neiges (proie alternative)- et qui subissent un accroissement des précipitations et des températures au travers des années. Nous avons utilisé une combinaison de modèles illustrant les liens causaux multiples entre les variables clés issues d'une base de données à long-terme (21 ans). Nous avons obtenu plusieurs scénarios possibles reliant les facteurs climatiques régionaux (Oscillation Arctique) et les températures et précipitations locales à la reproduction de nos 3 espèces. Nos résultats suggèrent que les facteurs climatiques régionaux et locaux présentent des impacts directs et indirects sur la reproduction du renard arctique et de l'oie des neiges. Le climat local présente un lien causal positif avec le succès de nidification de l'oie, alors que le climat local et régional démontrent un effet contrasté sur la proportion de renard en reproduction. Aucune relation entre les facteurs climatiques et l'abondance des lemmings n'a été trouvée. Nous avons observé des liens causaux positifs entre la reproduction du lemming, du renard et de l'oie, mettant en évidence les réponses numériques et fonctionnelles du renard arctique face à la variabilité de l'abondance de lemming. Notre étude est une des premières à mesurer les liens causaux et les forces d'interaction entre les espèces partageant un même réseau trophique, quantifiant ainsi la réponse numérique du prédateur et les interactions apparentes entre ses proies principales. Ces résultats améliorent notre compréhension des effets complexes du climat sur les interactions prédateur-proie et notre capacité à anticiper la réponse des réseaux trophiques aux changements climatiques en cours.

The arctic fox variant of the rabies virus (RABV) is enzootic in the circumpolar north. Reports of abortive RABV exposures motivated a retrospective analysis of sera from 41 arctic foxes (Vulpes lagopus) captured at Karrak Lake in Nunavut, Canada, during 2011-15. Estimated RABV antibody prevalence among foxes was 15% (95% confidence interval, 7-28%).

Predators largely affect ecosystems through trophic interactions, but they also can have indirect effects by altering nutrient dynamics and acting as ecosystem engineers. Arctic foxes (Vulpes lagopus) are ecosystem engineers that concentrate nutrients on their dens, creating biogeochemical hotspots with lush vegetation on the nutrient-limited tundra. Red foxes (V. vulpes) similarly engineer subarctic environments through their denning behavior, and have recently expanded onto the tundra where they now often occupy historical Arctic fox dens. We evaluated the impact of fox denning activity on the spatial behavior of other tundra wildlife by comparing predator and herbivore visits to 12 natal dens and adjacent control sites over two years using camera traps in northeastern Manitoba, where both fox species are sympatric. Both the capture rates and species richness of wildlife were significantly greater at fox dens relative to control sites. Predators were detected almost exclusively on dens occupied by foxes, where they were observed investigating and scavenging prey remains (carrion, feathers), suggesting carcass presence or fox presence attracts predators to den sites. Caribou (Rangifer tarandus) also visited dens more often than control sites, likely attracted by the enhanced vegetation typically found on dens. Our results suggest fox ecosystem engineering affects the spatial distribution of herbivores by enriching vegetation at dens, and other predators by providing carrion. Understanding how predators affect other organisms via non-trophic interactions provides an enriched view of their functional roles within ecosystems.

Rabies occurs throughout the Arctic, representing an ongoing public health concern for residents of northern communities. The Arctic fox (Vulpes lagopus) is the main reservoir of the Arctic rabies virus variant, yet little is known about the epidemiology of Arctic rabies, such as the ecological mechanisms driving where and when epizootics in fox populations occur. In this study, we provide the first portrait of the spatio-temporal spread of rabies across northern Canada. We also explore the impact of seasonal and multiannual dynamics in Arctic fox populations and climatic factors on rabies transmission dynamics. We analysed data on rabies cases collected through passive surveillance systems in the Yukon, Northwest Territories, Nunavut, Nunavik and Labrador from 1953 to 2014. In addition, we analysed a large and unique database of trapped foxes tested for rabies in the Northwest Territories and Nunavut from 1974 to 1984 as part of active surveillance studies. Rabies cases occurred in all Arctic regions of Canada and were relatively synchronous among foxes and dogs (Canis familiaris). This study highlights the spread of Arctic rabies virus variant across northern Canada, with contrasting rabies dynamics between different yet connected areas. Population fluctuations of Arctic fox populations could drive rabies transmission dynamics in a complex way across northern Canada. Furthermore, this study suggests different impacts of climate and sea ice cover on the onset of rabies epizootics in northern Canada. These results lay the groundwork for the development of epidemiological models to better predict the spatio-temporal dynamics of rabies occurrence in both wild and domestic carnivores, leading to better estimates of human exposure and transmission risk.

Many species of seabirds and waterfowl are predominantly colonial breeders, suggesting that colonial breeding is beneficial in some way. For Arctic nesting geese, colonial breeding may be an adaptive approach to minimize the risk of nest failure through predator swamping. Yet evidence for predator swamping is inconclusive. Species in low-lying coastal zones may have to manage tradeoffs between historical (e.g., predation) and increased frequency of other threats (e.g., coastal flooding associated with storm events). As the climate continues to change, coastal obligates will be subject to rising sea levels and an increase in extreme weather events, which could exacerbate the tradeoff between flooding and predation risk. We studied the effect of nest density on nest success in the colonial nesting Black Brant (Branta bernicla nigricans) on the Tutakoke River Colony in western Alaska and explored how density effects change during years of heavy Arctic fox (Alopex lagopus) predation and coastal flooding associated with storm events. We classified the colony as containing 2 strata that differed in elevation and vegetation, and were separated by the Tutakoke River. The stratum north of the Tutakoke River was 0.25 m lower in elevation and subject to periodic tidal flooding, while the southern stratum never flooded. Nest density positively affected brant nest success in years with high levels of fox predation. Nest success was greatest at the highest density during Normal years (low levels of fox predation and no flooding; northern stratum nest success 0.81 ± 0.01, southern stratum nest success 0.81 ± 0.01) and lowest in areas of low density during Fox years (northern stratum nest success 0.38 ± 0.01, southern stratum nest success 0.31 ± 0.01). Nest success was routinely higher during Normal years at all densities compared to Flood and Fox years. Our results support the hypothesis that colonial nesting, at least in tundra habitats, may have developed as a defense mechanism to swamp mammalian predators.

We compiled present-day observations and survey data of the avian and mammal fauna in the Islands of Four Mountains, Alaska, which primarily consist of marine-adapted species, to provide insight into the prehistoric fauna found in the archaeological record. We documented 63 bird species, mainly seabirds (67%). The majority (71%) of the estimated number of breeding birds are ledge nesters, predominately murres, and most (68%) spend the majority of their lives in the offshore environment. We lack good estimates for burrow and crevice nesters, which were impacted by and are still recovering from a period of arctic fox farming in the twentieth century. Marine mammals, predominately Steller sea lion, harbor seal, and sea otter, primarily use accessible land as haul-outs and are found in low numbers compared with the rest of the Aleutians. Red fox, the only land mammal, presently only reside on Chuginadak Island. While the present-day fauna is similar to the fauna recorded in the archaeological record, we found no indication of nesting by ancient murrelets and northern fulmars in the Island of Four Mountains, whereas both species were present in midden sites on Carlisle and Chuginadak Islands, with the possibility of prehistoric breeding colonies on Carlisle Island.

The rapidly growing interest for nature-based recreational activities threaten biodiversity values and increases the disturbance caused to wildlife. Several studies have demonstrated spatial and temporal activity shifts of animals in response to human disturbance. However, most studies investigate effects on how animals use an area of high tourism disturbance, such as a tourist resort or hiking trail, and not the effects of human disturbance at a key site for the animal, such as a denning or breeding site. In this study, we use photos from remote camera monitoring of arctic fox (Vulpes lagopus) dens with the aim to investigate the effects of various levels of tourism disturbance on the diurnal activity pattern and vigilance of breeding adult arctic foxes at the den site. We find a temporal shift towards a higher daytime activity at the den in response to high intensity tourism (71 ± 3.9% in disturbed areas compared with 53 ± 6.2% in undisturbed areas), which stands in contrast to an increased nocturnality seen in studies of many other species, including another study of arctic foxes. We suggest that the difference could be explained by the higher cost of avoiding a key site for the animal, as in this study, compared with avoiding a more general human disturbance in an area within the distribution range of the animal, as in most other studies. Increased time spent avoiding the perceived threat of humans could compromise other important activities and have potential negative effects on e.g. hunting or provisioning for the juveniles. Human disturbance focused at a key site, such as a denning or breeding site, can thereby be expected to have larger consequences than what is observed in most studies of disturbance effects on wildlife. Based on observations of both tourists and arctic foxes during close encounters in an area of high intensity tourism, we also find that they both respond behaviorally to each other. The potential for positive or negative feedback mechanisms in such relationships between tourists and wildlife highlights the importance of considering both sides of the complex interaction to find a balance between preserving biodiversity and ensuring continued possibilities for recreation.

Oil and gas development on the Arctic Coastal Plain (ACP) of Alaska, USA may have effects on Arctic-nesting birds. To estimate effects of industrial activity and investigator disturbance on avian productivity, we monitored nests of greater white-fronted geese (Anser albifrons) with digital cameras and periodic nest visits during 2013–2014 at 2 sites on the ACP. A disturbed site was adjacent to human-made infrastructure and industrial clean-up activities initiated at the onset of the study and a control site was >2 km from sources of industrial disturbance. We assessed variation in estimates of incubation constancy, nest survival, and predator behavior relative to site, year, and distance from industrial activity using nest photographs obtained at 1-minute intervals. We compared analysis of hourly nest survival informed by intensive monitoring with cameras to analysis of daily nest survival informed by traditional nest visit data obtained at intervals of 5–7 days to assess how method and time scale of sampling affect ecological inference. Geese in both sites exhibited high levels of nest attendance and initiated incubation breaks less than once per day. Observer-caused incubation breaks associated with nest visits (x¯ = 37.8 min) were longer than other types of incubation breaks (x¯ = 8.7 min), demonstrating a differential response by nesting geese to direct human encroachment versus indirect vehicular and aircraft traffic. During both years, geese were absent from nests more frequently in the disturbed (x¯ = 0.9 breaks/day) than control (x¯ = 0.6 breaks/day) site, and this break frequency was slightly higher for nests closer to industrial activity. In the year with high rates of depredation, nest survival was positively related to distance from industrial activity and abandoned infrastructure, consistent with predictions of industry-caused effects. This relationship, however, was not evident in the year with reduced predation pressure, likely because of annual variation in arctic fox (Vulpes lagopus) behavior. Analysis of nest survival probability informed by camera data allowed for detection of detailed patterns of variation that were not supported when using only visit data for the same nests. Observer visits were responsible for reductions of 7–35% in nest survival probability, highlighting the importance of minimizing, and controlling for, observer effects in studies of avian productivity. Indirect vehicular and aircraft disturbance posed less risk to nest survival than direct encroachment by observers at nest sites. Therefore, effects of industrial activities on avian productivity in the Arctic can be minimized through practices that limit direct encounters with nests. © 2017 The Wildlife Society.

Global climate change induces species range shifts and population expansion to higher latitudes in response to rising temperatures. One consequence of climate-induced range shifts is an increased sympatry between related but previously isolated species, potentially resulting in interspecific interactions and hybridization. The Arctic is more rapidly affected by climate warming than any region on Earth and resident species may be prone to interspecific hybridization due to the immigration of new colonizing species. The red fox Vulpes vulpes expanded its range into the Arctic during the twentieth century and is now in sympatry with the native Arctic fox Vulpes lagopus in many places. In this context, an Arctic-like fox displaying an unusual reddish winter coat was observed in 2013 in Nunavut, Canada, a phenotype unknown by Inuit people to date. In this study, we assessed the biological origin of this specimen with a multigenic approach using markers located on mitochondrial DNA, sex (X and Y) chromosomes, and autosomes (microsatellites and the melanocortin 1 receptor (MC1R) gene; a gene involved in the color polymorphism in canids). Our comparative analyses with genetic material from Arctic and red foxes displaying “classic” phenotypes (N = 14 and 16, respectively) suggested a pure Arctic fox origin of the specimen. Specifically, this fox yielded mtDNA, X-linked, Y-linked, and MC1R alleles specific to Arctic fox. Bayesian genetic assignment based on 16 microsatellite loci assigned it to Arctic fox with high confidence (q = 99.7%). Thus, a recent hybrid origin for this specimen is excluded. This alternative winter coat color polymorphism in Arctic fox requires further analyses to determine its underlying genetic mechanism.

Island subspecies of the Arctic fox Vulpes lagopus differ morphologically from the mainland subspecies. In particular, differences in cranial form may reflect varied biomechanical adaptations associated with hunting and feeding behaviours. We tested the hypothesis that the observed cranial differences between two island foxes (living on two North Pacific islands) and those living on the mainland have no impact on biomechanical performance during simulated biting. 3D cranial models of three Arctic fox subspecies were compared based on biomechanical parameters (e.g. local strain and large-scale deformation). Finite elements (FE) analyses were used to simulate equivalent biting loads, and geometric morphometrics was used to compare the modes of deformation among the models. The results showed differences in local strains and modes of global deformation among the three subspecies; the mainland subspecies was particularly distinct from the island subspecies. The representative cranium of the mainland subspecies experienced higher strain than that of both island subspecies during all bites. However, the findings highlight issues that arise when relating biomechanical performance, measured via FE analyses, to the foods consumed rather than to the mechanical properties of the individual’s diet. Additional physical properties data for each prey type are necessary to determine the extent to which the present findings reflect biomechanical adaptations to diet and prey acquisition.

Reproductive experience affects juvenile survival in a wide range of species with possible links to differences in foraging capacity and predation. Using supplementary feeding, we aimed to limit direct effect of prey abundance to investigate indirect effects of small-rodent availability and maternal experience on juvenile summer survival rates in an endangered population of arctic fox (Vulpes lagopus (L., 1758)). We used data spanning 7 years, included a complete small-rodent cycle, comprising 49 litters and 394 cubs. The effect of small-rodent abundance on juvenile survival depended on maternal breeding experience. Cubs born by first-time-breeding females had lower survival rate when small-rodent abundance was low compared with juveniles born to experienced mothers who remained unaffected. It was unlikely due to starvation, as physical condition was unrelated to survival. Instead, we favour the explanation that intraguild predation was an important cause of mortality. There was a negative relationship between survival and amount of time cubs were left unattended, suggesting that parental behaviour affected predation. We propose that a prey switch related to small-rodent abundance caused fluctuations in intraguild predation pressure and that inexperienced females were less able to cope with predation when small rodents were scarce.

Arctic Alaska is an important breeding ground for many migratory bird populations. A variety of factors associated with industrial development may impact nesting birds in this region, including increased nest predator populations associated with anthropogenic nesting and perching sites and the availability of anthropogenic food sources. We tested the indirect impact of oil development on nest survivorship in an artificial nest experiment at Prudhoe Bay, Alaska, 2012–2014, by establishing and monitoring 268 artificial shorebird nests and 221 artificial waterfowl nests and through monitoring of real shorebird nests (n = 186). Distance to infrastructure and roads, and area of infrastructure within 2 km2 of the nest did not significantly affect nest survival of artificial or real nests. Artificial nest survival was higher at shorebird than waterfowl nests. Cameras deployed at a subset of artificial shorebird nests documented nest predation by Arctic fox (Alopex lagopus), red fox (Vulpes vulpes), parasitic jaegers (Stercorarius parasiticus), pomarine jaegers (Stercorarius pomarinus), long-tailed jaegers (Stercorarius longicaudus), northern harriers (Circus cyaneus), and glaucous gulls (Larus hyperboreus). The presence of a camera had a positive effect on artificial shorebird nest survival, possibly due to cameras being placed on progressively older nests throughout the season. In conclusion, we did not detect an effect from infrastructure on nest survival at the scale of the study, in either real or artificial nests. We suggest caution when using artificial nests for Arctic research, given differences in survival of real and artificial nests in this study, and potential differences in nest predators.

Grazing lawns, patches of grazing‐tolerant plants with high nutrient value, provide important habitat for herbivores, and changes in abundance can impact herbivore populations. Grazing lawns are maintained in quality and quantity by repeated grazing and are a result of a positive feedback since the availability of grazing lawn can increase herbivore populations and increased herbivore populations can result in an increase in grazing lawn extent. We sampled aerial imagery from a long‐term dataset (>20 yr) at an internationally important breeding area for avian herbivores to model changes in grazing lawn abundance over time and identify the possible factors impacting those changes, including numbers of breeding birds, their primary predator, and spring phenology. Our data suggest that avian herbivores and their predators likely exert strong impacts on plant communities and may drive vegetation abundance. Decreases in the number of herbivore nests in our study coincided with an almost complete lack of grazing lawn in the mid‐2000s. Any factors dictating the amount of grazing lawn available for avian herbivores could strongly influence breeding success and the maximum size of these populations. Our results demonstrate the importance of studying complex interactions among predators, herbivores, and plants, and population moderation by both bottom‐up and top‐down processes.

Since the last Arctic Monitoring and Assessment Programme (AMAP) effort to review biological effects of the exposure to organohalogen compounds (OHCs) in Arctic biota, there has been a considerable number of new Arctic effect studies. Here, we provide an update on the state of the knowledge of OHC, and also include mercury, exposure and/or associated effects in key Arctic marine and terrestrial mammal and bird species as well as in fish by reviewing the literature published since the last AMAP assessment in 2010. We aimed at updating the knowledge of how single but also combined health effects are or can be associated to the exposure to single compounds or mixtures of OHCs. We also focussed on assessing both potential individual as well as population health impacts using population-specific exposure data post 2000. We have identified quantifiable effects on vitamin metabolism, immune functioning, thyroid and steroid hormone balances, oxidative stress, tissue pathology, and reproduction. As with the previous assessment, a wealth of documentation is available for biological effects in marine mammals and seabirds, and sentinel species such as the sledge dog and Arctic fox, but information for terrestrial vertebrates and fish remain scarce. While hormones and vitamins are thoroughly studied, oxidative stress, immunotoxic and reproductive effects need further investigation. Depending on the species and population, some OHCs and mercury tissue contaminant burdens post 2000 were observed to be high enough to exceed putative risk threshold levels that have been previously estimated for non-target species or populations outside the Arctic. In this assessment, we made use of risk quotient calculations to summarize the cumulative effects of different OHC classes and mercury for which critical body burdens can be estimated for wildlife across the Arctic. As our ultimate goal is to better predict or estimate the effects of OHCs and mercury in Arctic wildlife at the individual, population and ecosystem level, there remain numerous knowledge gaps on the biological effects of exposure in Arctic biota. These knowledge gaps include the establishment of concentration thresholds for individual compounds as well as for realistic cocktail mixtures that in fact indicate biologically relevant, and not statistically determined, health effects for specific species and subpopulations. Finally, we provide future perspectives on understanding Arctic wildlife health using new in vivo, in vitro, and in silico techniques, and provide case studies on multiple stressors to show that future assessments would benefit from significant efforts to integrate human health, wildlife ecology and retrospective and forecasting aspects into assessing the biological effects of OHC and mercury exposure in Arctic wildlife and fish. © 2019 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).

It is well known that competition, predation and fluctuating food resources can have strong effect on individual fitness and population dynamics. The complexity of natural systems can make it complicate to disentangle those processes, but environments with relatively simple food webs, and strong cyclic population dynamics offer contrasting conditions resembling experimental treatments. This thesis concerns the spatial and temporal implications of fluctuations in small rodent abundance on two trophic levels in a highly cyclic ecosystem, the Scandinavian mountain tundra. The first two chapters focus on plant biomass and spatiotemporal distribution in the Norwegian lemming (Lemmus lemmus), while the three last papers focus on the direct and indirect effects of small rodent fluctuations and territory quality on reproductive success, juvenile survival and group living in a lemming specialist mesopredator, the arctic fox (Vulpes lagopus). By developing, validating and applying a novel application of aerial photos for remote sensing of plant biomass (Chapter I), we found that food availability predicted lemming distribution during population peaks, but that they were more habitat specific during increase years when intraspecific competition was lower (Chapter II). Arctic fox reproduction is tightly connected to small rodent abundance but the effects of geographical variation in food availability is less well known. We used 17 years of population surveys of an arctic fox subpopulation in mid Sweden (Helagsfjällen) to investigate potential effects. During small rodent increase years, we found that arctic fox litter sizes were smaller in territories of intermediate plant productivity, compared to both more and less productive territories (Chapter III). This could be an effect of limited food availability together with increased presence of red foxes (Vulpes vulpes), a stronger and potentially lethal competitor. However, when small rodents peaked, and competition would be expected to decrease, we saw no effect of territory productivity. Based on a smaller data set concerning juvenile summer survival, we found that the mortality rate among juveniles born by first time breeding arctic fox females were more sensitive to low small rodent prey abundance (Chapter IV). We explain it with an increased predation pressure from top-predators that switch from small rodents to alternative prey when small rodents decline, as suggested by an observed positive effect on juvenile survival by adult presence on den sites. Arctic foxes are socially flexible, and several adults can share a den with the resident pair, potentially increasing juvenile survival and help in territorial defence. Returning to the 17-year data set, we tested the Resource Dispersion Hypothesis predicting that increased resource availability should increase group size (Chapter V). We found support for this prediction as group living increased during the small rodent peak phase. However, it remained unexpectedly high during the decrease phase, when resources are scarce. This could however be related to increased predation pressure, and an increasing benefit of group living. At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 1: Manuscript. Paper 3: Manuscript. Paper 5: Manuscript.

This study presents a complex approach for modeling the natural evolution of a population in terms of population number and dynamics of the genetic structure. A set of dynamic models that consider various types of natural selection was applied to describe possible mechanisms underlying the formation of existing genetic variations in litter sizes in coastal, inland, and farmed arctic fox populations (Alopex lagopus, family Canidae, order Carnivora). The r–K selection model for uniform population and the models with natural selection were assessed on various life cycle stages in a two-age population. The life cycle of arctic fox was fitted to the population model with two age stages. The different reproductive potentials and survivability of progeny on the early stage of life cycle were genetically determined using the model with a single diallelic gene. A monomorphism was obtained for a considered characteristic in a population of coastal arctic fox with constant food supply. Meanwhile, a polymorphism with cyclic fluctuations in population number and gene frequency was obtained in inland arctic fox populations, which could be due to cyclic fluctuations of prey. In farmed fox populations, the considered gene becomes pleiotropic (defines the survival rate of individuals on early and late stages of the life cycle) because of artificial selection performed by farmers to increase the reproductive success of breeders. The application of an appropriate model (with selection by pleiotropic gene) can be used to determine the elimination rate of low litter size alleles from the farmed populations. The possible applications of the proposed models for formulating and solving optimal control tasks in arctic fox populations are discussed too.

Creating farms for sable breeding was associated with the commercial destruction of natural populations and, consequently, the overall decline in the species number. The gene pool of the first farm-bred sable population in Russia, established in the vicinity of Moscow (“Pushkinskiy” fur farm), was formed by crossing of animals removed from nine natural populations. In the first eight years of farm operation, approximately one thousand animals were used for sable breeding; some of these animals were able to adapt to the farm management and, subsequently, to the selection for a number of quantitative traits in the period of industrial domestication. It took about ten years for breeders to work out the breeding and selection technologies, which became successfully employed in the established affiliated sable breeding farms. The main achievement in sable breeding over the 85-year historical period of breeding in Russia is the creation of two unique breeds, black sable (1969) and Saltykovskaya 1 (2007). In general, industrial domestication in fur farming and the subsequent breeding works made the fur of many species (mink, fox, Arctic fox) obtained from natural populations uncompetitive, which undoubtedly reduced the hunting interest in the animals living in the wild. Consequently, hunting for fur-bearing animals of most species decreased and has only local importance. Owing to the specific features of sable biology, the fur of farm-bred animals cannot yet completely replace the furs obtained by hunting; however, the farm-bred sable population is constantly growing. This review presents the results of the analysis of the level of genetic variability in natural and farm populations at nuclear and mitochondrial loci. The comparative analysis makes it possible to estimate the loss of genetic diversity upon the species adaptation to the new conditions of existence.

In territorial species, observed density dependence is often manifest in lowered reproductive output at high population density where individuals have fewer resources or are forced to inhabit low-quality territories. The Arctic fox (Vulpes lagopus) in Iceland is territorial throughout the year and feeds mostly on birds, since lemmings are absent from the country. Thus, the population does not exhibit short-term population cycles that are evident in most of the species' geographical range. The population has, however, gone through a major long-term fluctuation in population size. Because of the stability in hunting effort and reliable hunting records since 1958, the total number of adult foxes killed annually can be used as an index of population size (N t ). An index of carrying capacity (K) from population growth data for five separate time blocks during 1958-2007 revealed considerable variation in K and allowed a novel definition of population density in terms of K, or N t /K. Correlation analysis suggested that the reproductive rate was largely determined by the proportion of territorial foxes in the population. Variation in litter size and cub mortality was, on the other hand, related to climatic variation. Thus, Arctic foxes in Iceland engage in typical contest competition but can adapt their territory sizes in response to both temporal and spatial variation in carrying capacity, resulting in surprisingly little variation in litter size.

We studied the eff ect of the two environmental indices, the sub-polar gyre (SPG), and winter and summer North Atlantic Oscillation (NAO), together with mean annual winter and summer temperatures and geographic location on mandible size and body mass of Arctic foxes in Iceland (6345 and 2732 specimens, respectively) during the year of their death. We predicted that when favorable conditions prevailed, large specimens would be selected for, and vice versa. Body size and body mass were signifi cantly aff ected by the environmental parameters (i.e. SPG, NAO, ambient temperature and cloud cover) prevailing during the year of death. Th e eff ect of environmental conditions on body size was much stronger in the less productive region of eastern Iceland, apparently because in areas where food availability is meager, even a small diff erence in climate may tilt the balance from food suffi ciency to food shortage. Western Iceland comprises only a quarter of the total surface area of the country, but its productive seashores are twice as long as those of all the rest of the country combined. It is interesting to note that the eff ect of the SPG, a marine phenomenon in the oceans surrounding Iceland, is refl ected in the condition of the foxes more than the other climatic variables we used in this study, which are largely land-related. Because Arctic foxes in Iceland feed largely on marine birds and invertebrates, the SPG seems to encompass more accurate information regarding the direct ocean forces that aff ect food availability to the foxes.

This paper considers the integrated approach to modeling the dynamics of genetic structure and the number of natural population. A set of dynamic models with different types of natural selection is used to describe a possible mechanism for the fixing of a genetic diversity in size of the litter in coastal, continental and farmed populations of arctic fox (Alopex lagopus, Canidae, Carnivora) observed now. The most interesting results have been obtained with the model of population consisting of two stages of development. At that with the frame of this model a dynamics of population genetic structure on genotypes was analyzed to consider different reproductive abilities and fitnesses of pups on the early stage of lifecycle which defined by the single diallelic gene. This model allows to receive a monomorphism for coastal populations of arctic fox, where food resources are practically constant. As well the model allows polymorphism with cyclical fluctuations in the number and frequency of the gene in the continental populations due to regular fluctuating of rodent number, the major component of its food. In farmed populations by selective selection carried out by farmers to increase the reproductive success, this gene is a pleiotropic one (i. e., determining the survival rate of individuals both early and late stages of their life cycle); so an application of appropriate model (with the selection of pleiotropic gene) allows to get an adequate rate of elimination for small litters allele.

Arctic foxes Vulpes lagopus (L.) display a sharp 3- to 5-year fluctuation in population size where lemmings are their main prey. In areas devoid of lemmings, such as Iceland, they do not experience short-term fluctuations. This study focusses on the population dynamics of the arctic fox in Iceland and how it is shaped by its main prey populations. Hunting statistics from 1958-2003 show that the population size of the arctic fox was at a maximum in the 1950s, declined to a minimum in the 1970s, and increased steadily until 2003. Analysis of the arctic fox population size and their prey populations suggests that fox numbers were limited by rock ptarmigan numbers during the decline period. The recovery of the arctic fox population was traced mostly to an increase in goose populations, and favourable climatic conditions as reflected by the Subpolar Gyre. These results underscore the flexibility of a generalist predator and its responses to shifting food resources and climate changes.