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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

1. Habitat fragmentation may affect species distributions through, for example, altered resource availability and shifts in species interactions. Fragmentation by roads has had negative impacts on Fennoscandian alpine ecosystems, with reduction of habitats and connectivity for alpine species. Concurrently, infrastructure development causes influx of subsidies through roadkills and litter, which may facilitate expansion of boreal scavenging species, such as the red fox Vulpes vulpes, which may intensify negative interactions with alpine species. Hence, understanding the impact of subsidies within marginal alpine areas is imperative for successful conservation and management of particularly vulnerable alpine species. 2. We used snow tracking and camera traps in three alpine tundra areas in Norway to investigate whether the presence of boreal scavengers was positively associated with highways during winter, and if this contrasted the pattern of a critically endangered alpine species, the Arctic fox Vulpes lagopus. In summer, artificial nests were used to assess whether predation risk was related to proximity to highways. 3. During winter, the occurrence of red foxes was higher close to highways and decreased with increasing distance to highways, while the arctic fox showed no discernible pattern. Red fox occurrence increased with the number of edible items of anthropogenic origin located along highways, whereas arctic fox occurrence decreased. 4. The overall predation risk of artificial nests during summer was high (>31.2%) and increased with proximity to the highway in the area with highest traffic volume. 5. Synthesis and applications. Highways crossing alpine areas may attract boreal scavengers, possibly connected to increased access to subsidies of anthropogenic origin. Litter and food waste dominated available subsidies along highways in our study, and prevailing mitigating measures directed at reducing roadkill and movement restrictions may not be applicable to reduce negative effects of littering. We recommend actions focusing on informational campaigns, improved garbage disposal facilities and routines, and imposing fines for littering, to reduce negative impacts on vulnerable species. This is likely needed to achieve goals of ‘no impact’ from the physical loss of habitats due to road development. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

Translocation of captive-bred animals has become a widespread conservation practice to counteract species extinctions. We analyse and discuss the apparent success and shortcomings of Arctic fox (Vulpes lagopus) reintroductions in alpine tundra areas of Norway. We followed the fate of 915 foxes between 2007 and 2020 and estimated the apparent survival and reproductive success of captive-bred and released Arctic foxes, compared to wild-born descendants. Relationship to abundance of small rodents, population size, and age were explored. Overall, apparent survival and probability of breeding were similar between captive-bred and wild-born foxes, positively linked to rodent abundance. For wild-born foxes, both breeding propensity and litter size declined with increasing fox population size. This could be a first sign of the limited capacity of single tundra patches to house self-subsistent populations. Thus, facilitating and maintaining connectivity among remnant and re-established Arctic fox populations, creating functional metapopulations, is essential for further improvement and longterm survival. Relying on the combined measures of supplementary feeding and red-fox (Vulpes vulpes) control, the Arctic fox captive-breeding and reintroduction programme has so far been highly successful. However, anthropogenic drivers facilitating red fox invasion into the Arctic fox habitat, along with climate driven irregularities and dampened small rodent cycles, could inhibit the establishment of a self-sustained population. A more holistic ecosystem approach and conservation measures to restore alpine fauna should be considered.

1. Large carnivores influence ecosystem dynamics in multiple ways, for example, by suppressing meso-carnivores and providing carrions for smaller scavengers. Loss of large carnivores is suggested to cause meso-carnivore increase and expansion. Moreover, competition between meso-carnivores may be modified by the presence of larger carnivores. In tundra ecosystems, the smallest meso-carnivore, the Arctic fox, has experienced regional declines, whereas its larger and competitively superior congener, the red fox, has increased, potentially due to changes in the abundance of apex predators. 2. We explored if variation in the occurrence of wolverine and golden eagle impacted the occurrence and co-occurrence of the Arctic fox and red fox in relation to varying abundances of small rodents within the Scandinavian tundra. 3. We applied multi-species occupancy models to an extensive wildlife camera dataset from 2011–2020 covering 98 sites. Daily detection/non-detection of each species per camera trap site and study period (late winter; March–May) was stacked across years, and species occupancy was related to small rodent abundance while accounting for time of the year and status of simulated carcass. 4. The Arctic fox was more likely to co-occur with the red fox when the wolverine was present and less likely to co-occur with the red fox when golden eagles were present and the wolverine was absent. Red foxes increased in occupancy when co-occurring with the larger predators. The Arctic fox responded more strongly to small rodent abundance than the red fox and co-occurred more often with the other species at carcasses when rodent abundance was low. 5. Our findings suggest that the interspecific interactions within this tundra predator guild appear to be surprisingly intricate, driven by facets of fear of predation, interspecific mediation and facilitation, and food resource dynamics. These dynamics of intraguild interactions may dictate where and when conservation actions targeted towards the Arctic fox should be implemented.

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.

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

Concern has been raised regarding red fox (Vulpes Vulpes) population increase and range expansion into alpine tundra, directly and indirectly enhanced by human activities, including carrion supply, and its negative impact on native fauna. In this study, we used cameras on bait stations and hunting remains to investigate how spatiotemporal patterns of red fox scavenging were influenced by abundance and accessibility of live prey, i.e., small rodent population cycles, snow depth, and primary productivity. We found contrasting patterns of scavenging between habitats during winter. In alpine areas, use of baits was highest post rodent peaks and when snow depth was low. This probably reflected relatively higher red fox abundance due to increased reproduction or migration of individuals from neighboring areas, possibly also enhanced by a diet shift. Contrastingly, red fox use of baits in the forest was highest during rodent low phase, and when snow was deep, indicating a higher dependency of carrion under these conditions. Scavenging patterns by red fox on the pulsed but predictable food resource from hunting remains in the autumn revealed no patterns throughout the rodent cycle. In this study, we showed that small rodent dynamics influenced red fox scavenging, at least in winter, but with contrasting patterns depending on environmental conditions. In marginal alpine areas, a numerical response to higher availability of rodents possible lead to the increase in bait visitation the proceeding winter, while in more productive forest areas, low availability of rodents induced a functional diet shift towards scavenging.

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

Scavengers can have strong impacts on food webs, and awareness of their role in ecosystems has increased during the last decades. In our study, we used baited camera traps to quantify the structure of the winter scavenger community in central Scandinavia across a forest–alpine continuum and assess how climatic conditions affected spatial patterns of species occurrences at baits. Canonical correspondence analysis revealed that the main habitat type (forest or alpine tundra) and snow depth was main determinants of the community structure. According to a joint species distribution model within the HMSC framework, species richness tended to be higher in forest than in alpine tundra habitat, but was only weakly associated with temperature and snow depth. However, we observed stronger and more diverse impacts of these covariates on individual species. Occurrence at baits by habitat generalists (red fox, golden eagle, and common raven) typically increased at low temperatures and high snow depth, probably due to increased energetic demands and lower abundance of natural prey in harsh winter conditions. On the contrary, occurrence at baits by forest specialists (e.g., Eurasian jay) tended to decrease in deep snow, which is possibly a consequence of reduced bait detectability and accessibility. In general, the influence of environmental covariates on species richness and occurrence at baits was lower in alpine tundra than in forests, and habitat generalists dominated the scavenger communities in both forest and alpine tundra. Following forecasted climate change, altered environmental conditions are likely to cause range expansion of boreal species and range contraction of typical alpine species such as the arctic fox. Our results suggest that altered snow conditions will possibly be a main driver of changes in species community structure., In our study, we used baited camera traps to quantify the structure of the winter scavenger community in central Scandinavia across a forest–alpine continuum and assess how climatic conditions affected spatial patterns of species occurrences. Following forecasted climate change, altered environmental conditions is likely to cause range expansion of boreal species and range contraction of typical alpine species. Our results suggest that altered snow conditions will be a main driver of change.

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.