Spatiotemporal patterns of red fox scavenging in forest and tundra: the influence of prey fluctuations and winter conditions

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. Red fox . Scavenging . Small rodents . Habitat . Climate . Camera trapspublishedVersio

Context-dependent demographic and genetic effects of translocation from a captive breeding project

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.publishedVersio

Rodent population dynamics affect seedling recruitment in alpine habitats

Questions: How do rodents with cyclic population dynamics affect seedling recruitment in alpine habitats? Does disturbance fromrodents have larger implications on seedling recruitment in some plant communities than in others? Location: Snowbeds and sheltered heaths in the low-alpine zone in areas of Børgefjell and Dovrefjell, Norway. Methods: We recorded seedling emergence, rodent activity and cover of mosses, lichens, litter and bare ground in 270 plots in snowbeds and sheltered heaths in a rodent population peak year and in the following low-density year. Results: Seedling recruitment was positively correlated with disturbances from lemmings and voles in both years. More seedlings emerged in the low-density year than in the year of the population peak. Snowbeds had higher seedling recruitment than the sheltered heaths, but both habitats were equally affected by disturbances fromrodents. Conclusions: Rodent activity created gaps and increased seedling emergence in these alpine plant communities, particularly in the year after the rodent peak, both in snowbeds and sheltered heath habitats. Our study therefore suggests that regeneration patterns in alpine vegetation are tightly linked to the population cycles of lemmings and voles,which peak in density at 3- to 5-yr intervals.publishedVersionPaid Open Acces

Spatial distribution in Norwegian lemming Lemmus lemmus in relation to the phase of the cycle

Competition between individuals of the same or different species affects spatial distribution of organisms at any given time. Consequently, a species geographical distribution is related to population dynamics through density-dependent processes. Small Arctic rodents are important prey species in many Arctic ecosystems. They commonly show large cyclic fluctuations in abundance offering a potential to investigate how landscape characteristics relates to density-dependent habitat selection. Based on long-term summer trapping data of the Norwegian lemming (Lemmus lemmus) in the Scandinavian Mountain tundra, we applied species distribution modeling to test if the effect of environmental variables on lemming distribution changed in relation to the lemming cycle. Lemmings were less habitat specific during the peak phase, as their distribution was only related to primary productivity. During the increase phase, however, lemming distribution was, in addition, associated with landscape characteristics such as hilly terrain and slopes that are less likely to get flooded. Lemming habitat use varied during the cycle, suggesting density-dependent changes in habitat selection that could be explained by intraspecific competition. We believe that the distribution patterns observed during the increase phase show a stronger ecological signal for habitat preference and that the less specific habitat use during the peak phase is a result of lemmings grazing themselves out of the best habitat as the population grows. Future research on lemming winter distribution would make it possible to investigate the year around strategies of habitat selection in lemmings and a better understanding of a fundamental actor in many Arctic ecosystems.publishedVersio

Scavenger community structure along an environmental gradient from boreal forest to alpine tundra in Scandinavia

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 cam-era 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 dis-tribution 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.publishedVersio

Scavenger community structure along an environmental gradient from boreal forest to alpine tundra in Scandinavia

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 cam-era 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 dis-tribution 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.publishedVersio

Predator co-occurrence in alpine and Arctic tundra in relation to fluctuating prey

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

Fecal glucocorticoid metabolites as an indicator of adrenocortical activity in Arctic foxes (Vulpes lagopus) and recommendations for future studies

Measuring fecal glucocorticoid metabolites (fGCMs) is a widely used, non-invasive method for studies of stress in vertebrates. To study physiological responses in wild Arctic foxes (Vulpes lagopus) to perceived stressors such as fluctuating food availability, occurrence of competitors and predators and disturbance from human activities, a species-specific physiological validation of a method to evaluate adrenocortical activity is needed. Here we used 15 captive Arctic foxes (both males and females and juveniles and adults) to investigate fGCM concentrations following ACTH injection (physiological validation), or handling alone and compared them with their respective baseline concentrations prior to the treatments. A 5α-pregnane-3ß,11ß,21-triol-20-one enzyme immunoassay measured significant fGCM increases following both treatments. The time lags to reach peak fGCM values were 9.3 ± 1.3 h and 12.8 ± 1.7 h for ACTH and handling treatment, respectively. Concentrations of fGCMs varied a lot between individuals, but not attributed to sex nor age of the foxes. However, we found a negative relationship between boldness and fGCM concentrations. Faecal glucocorticoid metabolites concentrations did not change significantly over a period of 48 h in samples kept at temperatures reflecting winter and summer means. This would allow the collection of samples up to two days old in the wild regardless of the season. We conclude that our successfully validated method for measuring fGCMs can be used as a non-invasive tool for studies exploring various stressors both in wild and captive Arctic foxes.publishedVersio

Conservation of the endangered Arctic fox in Norway - are successful reintroductions enough?

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.publishedVersio

Predation of endangered Arctic foxes by Golden eagles: What do we know?

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