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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.

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.

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.

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

This is the pre-peer reviewed version of the following article: Urbani, Tryland, Ehrich, Fuglei, Battilani, Balboni. Ancient origin and genetic segregation of canine circovirus infecting arctic foxes (Vulpes lagopus) in Svalbard and red foxes (Vulpes vulpes) in Northern Norway. Transboundary and Emerging Diseases. 2020;00:1-11, which has been published in final form at https://doi.org/10.1111/tbed.13783. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Use of Self-Archived Versions. Canine circovirus (CanineCV) is a relatively new viral species, belonging to the family Circoviridae, whose pathogenic role is still uncertain. Since its first description in one domestic dog in 2011 from the USA, several reports have been documenting its distribution worldwide. Recently, CanineCV was also detected in wild animals such as wolves, foxes and badgers. In order to investigate the presence and the genetic characteristics of CanineCV in foxes of Arctic and Sub‐Arctic regions, the presence of CanineCV DNA in internal organs (liver and spleen) of 51 arctic foxes (Vulpes lagopus) from Svalbard archipelago and 59 red foxes (Vulpes vulpes) from Northern Norway, sampled from 1996 to 2001 and from 2014 to 2018, respectively, was screened by real‐time PCR. CanineCV was detected in 11/51 arctic foxes and in 10/59 red foxes, backdating the circulation of the virus at least to 1996 in the arctic fox population. The complete genome of 14 identified CanineCV was sequenced and analysed showing an identity higher than 80.8% with the reference strains available to date. According to the species demarcation threshold of 80% genome‐wide nucleotide sequence identity for members of the family Circoviridae provided by International Committee on Taxonomy of Viruses (ICTV), all the CanineCV belong to a single species. Phylogenetic analysis revealed that all the CanineCV were subdivided into five main clusters with one including only CanineCV identified in foxes. Furthermore, CanineCV identified in arctic foxes and red foxes formed two distinct lineages. From these data, we hypothesize that the viral transmission did not occur between the two species of foxes as a consequence of the lack of contact between the two hosts or that the virus acquired mutations in the time elapsed between the samplings.

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.

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.

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.

Predators can affect prey through mechanisms other than predation; for example, redistributing resources could modify habitats favorably for other organisms. We examined Arctic fox (Vulpes lagopus) den use by lemmings, their primary prey, in winter during a year of low lemming densities. We found winter nests, which are built by lemmings under snow, on 69% of fox dens, whereas no control sites had nests. In August, dens had twice the vegetation cover and 50% greater nitrogen content in grass than controls, suggesting soil enrichment by foxes increased food quantity and quality for herbivores. Snow was ~4 times thicker in April on dens than controls, and 1.4 times thicker on dens with lemming nests than dens without, suggesting lemmings choose thicker snow for thermal insulation. Snow cover thickness was positively related to vegetation cover on dens, but not on control sites. Thus, Arctic foxes not only prey on lemmings but also engineer productive habitat that attracts lemmings. During winters with low lemming densities, when foxes often leave the denning area and predation risk is lower, fox dens may provide a refuge that could buffer the effects of deteriorating snow conditions with Arctic warming. This additional mechanism of predators interacting with their prey illustrates how ecosystem engineers potentially alter food web interactions and highlights the importance of integrating these bodies of theory in attempts to understand community dynamics.

The dominant white coat colour of farmed blue fox is inherited as a monogenic autosomal dominant trait and is suggested to be embryonic lethal in the homozygous state. In this study, the transcripts of KIT were identified by RT-PCR for a dominant white fox and a normal blue fox. Sequence analysis showed that the KIT transcript in normal blue fox contained the full-length coding sequence of 2919 bp (GenBank Acc. No KF530833), but in the dominant white individual, a truncated isoform lacking the entire exon 12 specifically co-expressed with the normal transcript. Genomic DNA sequencing revealed that a single nucleotide polymorphism (c.1867+1G>T) in intron 12 appeared only in the dominant white individuals and a 1-bp ins/del polymorphism in the same intron showed in individuals representing two different coat colours. Genotyping results of the SNP with PCR-RFLP in 185 individuals showed all 90 normal blue foxes were homozygous for the G allele, and all dominant white individuals were heterozygous. Due to the truncated protein with a deletion of 35 amino acids and an amino acid replacement (p.Pro623Ala) located in the conserved ATP binding domain, we propose that the mutant receptor had absent tyrosine kinase activity. These findings reveal that the base substitution at the first nucleotide of intron 12 of KIT gene, resulting in skipping of exon 12, is a causative mutation responsible for the dominant white phenotype of blue fox.

We used data from aerial surveys (1992-2010) of >100,000 km(2) and ground surveys (1998-2004) of >150 km(2) to estimate the density and abundance of birds on the North Slope of Alaska (U.S.A.). In the ground surveys, we used double sampling to estimate detection ratios. We used the aerial survey data to compare densities of birds and Arctic fox (Vulpes lagopus), the major nest predator of birds, on the North Slope, in Prudhoe Bay, and in nearby areas. We partitioned the Prudhoe Bay oil field into 2 × 2 km plots and determined the relation between density of aquatic birds and density of roads, buildings, and other infrastructure in these plots. Abundance and density (birds per square kilometer) of 3 groups of aquatic birds-waterfowl, loons, and grebes; shorebirds; and gulls, terns, and jaegers-were highest in the National Petroleum Reserve-Alaska (NPRA) and lowest in the Arctic National Wildlife Refuge. Six other major wetlands occur in the Arctic regions of Canada and Russia, but the largest population of aquatic birds was in the NPRA. Aquatic birds were concentrated in the northern part of the NPRA. For example, an area that covered 18% of the NPRA included 53% of its aquatic birds. The aerial surveys showed that bird density was not lower and fox density was not higher in Prudhoe Bay than in surrounding areas. Density of infrastructure did not significantly affect bird density for any group of species. Our results establish that the NPRA is one of the most important areas for aquatic birds in the Arctic. Our results and those of others also indicate that oil production, as practiced in Prudhoe Bay, does not necessarily lead to substantial declines in bird density or productivity in or near the developed areas. Prioridades para la Conservacion de Aves en el Norte de Alaska.

Many Arctic animals carry high body burdens of organochlorine contaminants (OCs) as a result of long-range transport of persistent pollutants. It has been shown that seasonal mobilization of body fat in these species results in increased blood concentration of OCs. The authors investigated OC assimilation, tissue distribution, and biotransformation in farmed Arctic fox (Vulpes lagopus) continuously fed a diet containing contaminated minke whale blubber or lard (control) from 8 wk of age in August 2003, until sampling when they were at their fattest (in November 2004) and leanest (in June 2005). Markedly higher tissue (liver, adrenals, brain, and blood) OC levels were found in June than in November despite low exposure to OCs during emaciation, suggesting that OCs had been redistributed from adipose tissues to vital organs. There were no differences in the activities of hepatic biotransforming enzymes between exposed fat and control fat foxes, except for 16α-hydroxylation, which was higher in exposed fat foxes. In emaciated foxes, ethoxyresorufin activity was higher in exposed than in control foxes, indicating an enhanced potential for toxicity of OCs with emaciation. Lower activities of 6β- and 2β-hydroxylation were found in lean than in fat foxes, irrespective of OC treatment. The results show that emaciation increase the toxic potential of accumulated OCs and emphasize that body adiposity must be considered when time-trend analyses, risk assessments, and effect studies are designed. Environ Toxicol Chem 2013;32:1784–1792. © 2013 SETAC

The direct and interactive effects of climate change on host species and infectious disease dynamics are likely to initially manifest\ at latitudinal extremes. As such, Alaska represents a region in the United States for introspection on climate change and disease. Rabies is enzootic among arctic foxes (Vulpes lagopus) throughout the northern polar region. In Alaska, arctic and red foxes (Vulpes vulpes) are reservoirs for rabies, with most domestic animal and wildlife cases reported from northern and western coastal Alaska. Based on passive surveillance, a pronounced seasonal trend in rabid foxes occurs in Alaska, with a peak in winter and spring. This study describes climatic factors that may be associated with reported cyclic rabies occurrence. Based upon probabilistic modelling, a stronger seasonal effect in reported fox rabies cases appears at higher latitudes in Alaska, and rabies in arctic foxes appear disproportionately affected by climatic factors in comparison with red foxes. As temperatures continue a warming trend, a decrease in reported rabid arctic foxes may be expected. The overall epidemiology of rabies in Alaska is likely to shift to increased viral transmission among red foxes as the primary reservoir in the region. Information on fox and lemming demographics, in addition to enhanced rabies surveillance among foxes at finer geographic scales, will be critical to develop more comprehensive models for rabies virus transmission in the region.

Summary 1. More than a quarter of the world’s carnivores are threatened, often due to multiple and complex causes. Considerable research efforts are devoted to resolving the mechanisms behind these threats in order to provide a basis for relevant conservation actions. However, even when the underlying mechanisms are known, specific actions aimed at direct support for carnivores are difficult to implement and evaluate at efficient spatial and temporal scales. 2. We report on a 30-year inventory of the critically endangered Fennoscandian arctic fox Vulpes lagopus L., including yearly surveys of 600 fox dens covering 21 000 km 2 . These surveys showed that the population was close to extinction in 2000, with 40–60 adult animals left. However, the population subsequently showed a fourfold increase in size. 3. During this time period, conservation actions through supplementary feeding and predator removal were implemented in several regions across Scandinavia, encompassing 79% of the area. To evaluate these actions, we examined the effect of supplemental winter feeding and red fox control applied at different intensities in 10 regions. A path analysis indicated that 47% of the explained variation in population productivity could be attributed to lemming abundance, whereas winter feeding had a 29% effect and red fox control a 20% effect. 4. This confirms that arctic foxes are highly dependent on lemming population fluctuations but also shows that red foxes severely impact the viability of arctic foxes. This study also highlights the importance of implementing conservation actions on extensive spatial and temporal scales, with geographically dispersed actions to scientifically evaluate the effects. We note that population recovery was only seen in regions with a high intensity of management actions. 5. Synthesis and applications. The present study demonstrates that carnivore population declines may be reversed through extensive actions that target specific threats. Fennoscandian arctic fox is still endangered, due to low population connectivity and expected climate impacts on the distribution and dynamics of lemmings and red foxes. Climate warming is expected to contribute to both more irregular lemming dynamics and red fox appearance in tundra areas; however, the effects of climate change can be mitigated through intensive management actions such as supplemental feeding and red fox control.

We performed a systematic characterization of the hepatic vitamin A storage in mammals and birds of the Svalbard archipelago and Greenland. The liver of top predators, including polar bear, arctic fox, bearded seal, and glaucous gull, contained about 10–20 times more vitamin A than the liver of all other arctic animals studied, as well as their genetically related continental top predators. The values are also high compared to normal human and experimental animals like mouse and rat. This massive amount of hepatic vitamin A was located in large autofluorescent lipid droplets in hepatic stellate cells (HSCs; also called vitamin A-storing cells, lipocytes, interstitial cells, fat-storing cells, or Ito cells). The droplets made up most of the cells' cytoplasm. The development of such an efficient vitamin A-storing mechanism in HSCs may have contributed to the survival of top predators in the extreme environment of the arctic. These animals demonstrated no signs of hypervitaminosis A. We suggest that HSCs have capacity to take-up and store large amounts of vitamin A, which may play a pivotal role in maintenance of the food web, food chain, biodiversity, and eventually ecology of the arctic. Anat Rec, 2012. © 2012 Wiley Periodicals, Inc.

Summary 1. Flows of nutrients and energy across ecosystem boundaries have the potential to subsidize consumer populations and modify the dynamics of food webs, but how spatio-temporal variations in autochthonous and allochthonous resources affect consumers’ subsidization remains largely unexplored. 2. We studied spatio-temporal patterns in the allochthonous subsidization of a predator living in a relatively simple ecosystem. We worked on Bylot Island (Nunavut, Canada), where arctic foxes (Vulpes lagopus L.) feed preferentially on lemmings (Lemmus trimucronatus and Dicrostonyx groenlandicus Traill), and alternatively on colonial greater snow geese (Anser caerulescens atlanticus L.). Geese migrate annually from their wintering grounds (where they feed on farmlands and marshes) to the Canadian Arctic, thus generating a strong flow of nutrients and energy across ecosystem boundaries. 3. We examined the influence of spatial variations in availability of geese on the diet of fox cubs (2003–2005) and on fox reproductive output (1996–2005) during different phases of the lemming cycle. 4. Using stable isotope analysis and a simple statistical routine developed to analyse the outputs of a multisource mixing model (SIAR), we showed that the contribution of geese to the diet of arctic fox cubs decreased with distance from the goose colony. 5. The probability that a den was used for reproduction by foxes decreased with distance from the subsidized goose colony and increased with lemming abundance. When lemmings were highly abundant, the effect of distance from the colony disappeared. The goose colony thus generated a spatial patterning of reproduction probability of foxes, while the lemming cycle generated a strong temporal variation of reproduction probability of foxes. 6. This study shows how the input of energy owing to the large-scale migration of prey affects the functional and reproductive responses of an opportunistic consumer, and how this input is spatially and temporally modulated through the foraging behaviour of the consumer. Thus, perspectives of both landscape and foraging ecology are needed to fully resolve the effects of subsidies on animal demographic processes and population dynamics.

Summary 1. Input of external subsidies in the Arctic may have substantial effects on predator populations that otherwise would have been limited by low local primary productivity. 2. We explore life-history traits, age-specific fecundity, litter sizes and survival, and the population dynamics of an Arctic fox (Vulpes lagopus) population to explore the influence of the spatial distribution and temporal availability of its main prey; including both resident and migrating (external) prey resources. 3. This study reveals that highly predictable cross-boundary subsidies from the marine food web, acting through seasonal access to seabirds, sustain larger local Arctic fox populations. Arctic fox dens located close to the coast in Svalbard were found to have higher occupancy rates, as expected from both high availability and high temporal and spatial predictability of prey resources (temporally stable external subsidies). Whereas the occupancy rate of inland dens varied between years in relation to the abundance of reindeer carcasses (temporally varying resident prey). 4. With regard to demography, juvenile Arctic foxes in Svalbard have lower survival rates and a high age of first reproduction compared with other populations. We suggest this may be caused by a lack of unoccupied dens and a saturated population.

Aim How the ecological neighbourhoods of coast and forest affect arctic tundra ecosystems is a pressing question as the circumpolar tundra belt is shrinking under global warming. Mobile facultative scavengers are likely to negatively impact tundra biodiversity as dominant competitors or predators, if they spill over into tundra. Here, we provide the first quantitative assessments of the structure of a scavenger guild in low arctic tundra with emphasis on how it changes along spatial gradients from neighbouring ecosystems (i.e. forest and coast) and with altitude (i.e. productivity gradients). We also assess the likelihood of interactions between guild members that may negatively impact vulnerable tundra species. Location North-eastern part of Norway. Methods Extensive records of scavenger prevalence were obtained by deploying automatic digital cameras at experimental carcasses in tundra regions covering several thousand square kilometres and three winters in northern Norway. Main conclusions We found short-range neighbourhood effects of forest and coast within the tundra scavenger guild. Species richness declined steeply with decreasing distance from the neighbouring ecosystems, in particular subarctic forest, and with increasing altitude. Bird species with strongholds in forest (golden eagle Aquila chrysaetos and hooded crow Corvus cornix) or along the coast (white-tailed eagle Haliaeetus albicilla) were mostly responsible for short-range neighbourhood effects on guild structure. However, the two most abundant guild members, the common raven Corvus corax and the red fox Vulpes vulpes, exhibited no spatial patterns within the range of neighbourhoods and altitudes examined. There was a clear diurnal segregation in the use of carcasses between birds and mammals reducing the likelihood of direct interactions between these two taxa. Presence of red fox appeared to exclude the arctic fox Vulpes lagopus, the only endemic tundra species within the guild, from carcasses.

Summary 1. Most studies addressing the causes of the recent increases and expansions of mesopredators in many ecosystems have focused on the top-down, releasing effect of extinctions of large apex predators. However, in the case of the northward expansion of the red fox into the arctic tundra, a bottom-up effect of increased resource availability has been proposed, an effect that can counteract prey shortage in the low phase of the multi-annual rodent cycle. Resource subsidies both with marine and with terrestrial origins could potentially be involved. 2. During different phases of a multi-annual rodent cycle, we investigated the seasonal dynamics and spatial pattern of resource use by red foxes across a coast to inland low arctic tundra gradient, Varanger Peninsula, Norway. We employed two complementary methods of diet analyses: stomach contents and stable isotope analysis. 3. We found that inland red foxes primarily subsisted on reindeer carrions during the low phase of a small rodent population cycle. Lemmings became the most important food item towards the peak phase of the rodent cycle, despite being less abundant than sympatric voles. Isotopic signatures of tissue from both predator and prey also revealed that red foxes near the coast used marine-derived subsidies in the winter, but these allochthonous resources did not spillover to adult foxes living beyond 20–25 km from the coast. 4. Although more needs to be learned about the link between increasing primary productivity due to climatic warming and trophic dynamics in tundra ecosystems, we suggest that changes in reindeer management through a bottom-up effect, at least regionally, may have paved the way towards the establishment of a new mesopredator in the tundra biome.

Summary There are five genes encoding melanocortin receptors. Among canids, the genes have mainly been studied in the dog (MC1R, MC2R and MC4R). The MC4R gene has also been analysed in the red fox. In this report, we present a study of chromosome localization, comparative sequence analysis and polymorphism of the MC3R gene in the dog, red fox, arctic fox and Chinese raccoon dog. The gene was localized by FISH to the following chromosome: 24q24-25 in the dog, 14p16 in the red fox, 18q13 in the arctic fox and NPP4p15 in the Chinese raccoon dog. A high identity level of the MC3R gene sequences was observed among the species, ranging from 96.0% (red fox – Chinese raccoon dog) to 99.5% (red fox – arctic fox). Altogether, eight polymorphic sites were found in the red fox, six in the Chinese raccoon dog and two in the dog, while the arctic fox appeared to be monomorphic. In addition, association of several polymorphisms with body weight was analysed in red foxes (the number of genotyped animals ranged from 319 to 379). Two polymorphisms in the red fox, i.e. a silent substitution c.957A>C and c.*185C>T in the 3′-flanking sequence, showed a significant association (P

Movement is a prominent process shaping genetic population structure. In many northern mammal species, population structure is formed by geographic distance, geographical barriers and various ecological factors that influence movement over the landscape. The Arctic fox Vulpes lagopus is a highly mobile, opportunistic carnivore of the Arctic that occurs in two main ecotypes with different ecological adaptations. We assembled microsatellite data in 7 loci for 1834 Arctic foxes sampled across their entire distribution to describe the circumpolar population structure and test the impact of (1) geographic distance, (2) geographical barriers and (3) ecotype designation on the population structure. Both Structure and Geneland demonstrated distinctiveness of Iceland and Scandinavia whereas low differentiation was observed between North America-northern Greenland, Svalbard and Siberia. Genetic differentiation was significantly correlated to presence of sea ice on a global scale, but not to geographical distance or ecotype designation. However, among areas connected by sea ice, we recorded a pattern of isolation by distance. The maximum likelihood approach in Migrate suggested that connectivity across North America-northern Greenland and Svalbard was particularly high. Our results demonstrate the importance of sea ice for maintaining connectivity between Arctic fox populations and we therefore predict that climate change will increase genetic divergence among populations in the future.

One obvious threat to the endangered arctic fox Vulpes lagopus population in Fennoscandia is competition with the larger red fox Vulpes vulpes, which may have expanded its range towards the alpine tundra because of increased food availability in the low-alpine and subalpine region. The steady increase in the number of vacation cabins and roads, and thus also in human garbage and road-killed animals, may subsidise easily available food resources and improve red fox survival in these otherwise marginal areas. In Borgefjell National Park, Norway, 14 of 27 known arctic fox dens were used by arctic fox during 2001-2005. The dens that were used were situated at higher altitudes, farther from natural red fox habitats, than unused dens. In the best of our logistic regression models, there was also a statistical negative effect of the number of cabins within 7 × 7 km squares around the den sites. Hence, our results support the prediction that the arctic fox is less likely to use areas where human activity...

Anthropogenic fragmentation of habitat and populations is recognized as one of the most important factors influencing loss of biodiversity. Since it is difficult to quantify demographic parameters ...

In food webs heavily influenced by multi-annual population fluctuations of key herbivores, predator species may differ in their functional and numerical responses as well as their competitive ability. Focusing on red and arctic fox in tundra with cyclic populations of rodents as key prey, we develop a model to predict how population dynamics of a dominant and versatile predator (red fox) impacted long-term growth rate of a subdominant and less versatile predator (arctic fox). We compare three realistic scenarios of red fox performance: (1) a numerical response scenario where red fox acted as a resident rodent specialist exhibiting population cycles lagging one year after the rodent cycle, (2) an aggregative response scenario where red fox shifted between tundra and a nearby ecosystem (i.e. boreal forest) so as to track rodent peaks in tundra without delay, and (3) a constant subsidy scenario in which the red fox population was stabilized at the same mean density as in the other two scenarios. For all three scenarios it is assumed that the arctic fox responded numerically as a rodent specialist and that the mechanisms of competition is of a interference type for space, in which the arctic fox is excluded from the most resource rich patches in tundra. Arctic fox is impacted most by the constant subsidy scenario and least by the numerical response scenario. The differential effects of the scenarios stemmed from cyclic phase-dependent sensitivity to competition mediated by changes in temporal mean and variance of available prey to the subdominant predator. A general implication from our result is that external resource subsidies (prey or habitats), monopolized by the dominant competitor, can significantly reduce the likelihood for co-existence within the predator guild. In terms of conservation of vulnerable arctic fox populations this means that the likelihood of extinction increases with increasing amount of subsidies (e.g. carcasses of large herbivores or marine resources) in tundra and nearby forest areas, since it will act to both increase and stabilize populations of red fox.

Arctic regions are expected to experience pronounced changes in climate during the current century. Large numbers of waterfowl breed in these regions, and any climate induced changes are likely to have consequences for their demographics. Moreover, environmental changes experienced during migration and on the wintering grounds may also have impacts but remain poorly understood. We investigate the role of climate variation during breeding, migration and wintering, while controlling for possible effects of mammalian predation and density dependence on the reproduction of Svalbard breeding barnacle geese Branta leucopsis using 40 years of observations. Breeding success was significantly positively correlated with temperature on both the wintering grounds (Scotland) and breeding grounds (Svalbard), but negatively correlated with the number of days of strong cross-winds during the northward migration period. These factors remained significant when controlling for a strong negative effect of population size. Goose reproduction on Svalbard was also linked to fluctuations in arctic fox Alopex lagopus populations occurring elsewhere in the arctic. This reveals the importance of mammalian predation, which may vary as a non-linear function of conditions within the wider arctic region. Climate predictions were used to project barnacle goose reproduction and hence the population until 2050. These simulations suggest the population will grow at between 1% and 2.7% per year, in response to increasing temperatures. However, it is harder to predict how changes in other factors, such as reductions in sea ice, may impact on arctic breeding birds.

Animals are exposed to environmental factors that influence their life history and body size. Here we used the Arctic fox (Vulpes lagopus) as an indicator of the complex links between largescale environmental variables that influence both marine and tundra trophic dynamics to demonstrate how they affect the fox’s body size and abundance. The Arctic fox inhabits throughout Iceland, where it preys mainly on birds. We studied the effects of the Sub-Polar Gyre (SPG), winter and summer North Atlantic Oscillation (NAO), mean annual winter and summer temperature, and geographic sector (eastern and western Iceland, which differ in their ecology) on variations in mandible size (6345 specimens) and body mass (2732 specimens) as well as abundance on the Arctic fox in Iceland. We found that (a) SPG index negatively affected male mandible length as well as body mass of both sexes. SPG was also negatively related to fox abundance. (b) Summer NAO had a negative effect on Arctic foxes, that is, cold summers were correlated with shorter mandibles and lower body mass. (c) Winter NAO had a significant negative effect (although weaker than that of summer NAO) on female mandible length, but not on body mass. (d) Summer temperature had a positive effect on female mandible length, but no effect on body mass. However, winter temperature had no effect on either the mandible or body mass. (e) Foxes in the eastern sector had shorter mandibles and were of lighter mass than those in the western sector. We suggest that climate conditions during the growth period of the young affected their final size both directly, by influencing energy metabolism for maintenance, but mainly through their effects on food availability. As far as we are aware, this is the first report that the SPG has an effect on vertebrates, let alone terrestrial ones.

Comprehensive population parameter data are useful for assessing effective conservation actions. The Fennoscandian arctic fox Alopex lagopus is critically endangered and the population size is estimated at 120 individuals that are fragmented into four isolated populations. Here, we use molecular tracking and visual observations to estimate population size and survival in one of the populations on the Swedish mountain tundra during a year of low food availability. We collected 98 arctic fox faecal samples during the winter of 2006 and recorded visual observations of ear-tagged individuals during the summer of 2005 and 2006. The faecal samples were analysed for variation in nine microsatellite loci and matched to the genetic profiles of previously ear-tagged individuals from 2001 to 2005. During winter 2006, the minimum number alive was 12 individuals using visual observations, 30 using molecular tracking and 36 by combining the datasets. Population size was estimated through mark–recapture for the molecular tracking and visual observation datasets and through rarefaction analyses for molecular tracking data. The mark–recapture estimate for visual observations was uninformative due to the large confidence interval (CI) (i.e. 6–212 individuals). Based on the molecular tracking dataset combined with the minimum number alive for visual observations and molecular tracking, we concluded a consensus population size of 36–55 individuals. We also estimated the age-specific finite survival rate during 1 year (July 2005 to July 2006) by combining molecular tracking with visual observations. Juvenile survival on a yearly basis was 0.08 (95% CI 0.02–0.18) while adults had a survival of 0.59 (95% CI 0.39–0.82). Juveniles displayed a lower survival than the adults during autumn (P

Skjonghclleren is a marine-cut cave with 15–20. m thick pre-Holocene sediments. Corings and excavations reveal three beds of extremely fine-grained, laminated sediments alternating with blocky sediments. The laminated beds are interpreted as glaciolacustrine sediments deposited subglacially at times when ice sheets covered the area, suggesting at least three glaciations after the cave was formed. The blocky/diamictic sediments were formed by frost-shattered blocks from the roof of the cave during ice-free periods, and mixing with the fines through slow mass movements along the floor of the cave. In the diamictic sediment beneath the uppermost laminated bed, almost 7,000 bone and teeth fragments of birds, mammals and fish were found. Birds dominated, with little auk and brunnich's guillemot as the most frequently occurring species. Arctic fox was the dominating mammal. During climatic optimum of the interstadial, conditions seem to have been similar to present-day coastal Finnmark, with North Atlantic warm water entering the Norwegian Sea. Two radiocarbon dates on bones and three Uranium series dates on speleothems from this bed all cluster around 30,000 B.P., i.e., the end of the Alesund interstadial. Above the uppermost laminated bed, bone fragments of birds, fish and mammals, deposited between c. 12,000 and c. 10,000 B.P., were found. Little auk dominate. The occurrence of squirrel is worth noting since it is limited mainly to areas with coniferous forest today. The beds below the 30,000B.P. bed are poorly dated or undated, but it is tentatively concluded that the entire sediment sequence was deposited during the Weichselian stage. It seems that the cave was formed at a high relative sea-level stand sometime during the Early Weichsclian. Two recorded palaeomagnetic excursions seem to correlate with the Laschamp/Olby and the Lake Mungo events, respectively.

1. Many ecosystems are characterized by pulses of dramatically higher than normal levels of foods (pulsed resources) to which animals often respond by caching foods for future use. However, the extent to which animals use cached foods and how this varies in relation to fluctuations in other foods is poorly understood in most animals. 2. Arctic foxes Alopex lagopus (L.) cache thousands of eggs annually at large goose colonies where eggs are often superabundant during the nesting period by geese. We estimated the contribution of cached eggs to arctic fox diets in spring and autumn, when geese were not present in the study area, by comparing stable isotope ratios (delta(13)C and delta(15)N) of fox tissues with those of their foods using a multisource mixing model in Program IsoSource. 3. The contribution of cached eggs to arctic fox diets was inversely related to collared lemming Dicrostonyx groenlandicus (Traill) abundance; the contribution of cached eggs to overall fox diets increased from < 28% in years when collared lemmings were abundant to 30-74% in years when collared lemmings were scarce. 4. Further, arctic foxes used cached eggs well into the following spring (almost 1 year after eggs were acquired) - a pattern that differs from that of carnivores generally storing foods for only a few days before consumption. 5. This study showed that long-term use of eggs that were cached when geese were superabundant at the colony in summer varied with fluctuations in collared lemming abundance (a key component in arctic fox diets throughout most of their range) and suggests that cached eggs functioned as a buffer when collared lemmings were scarce.

Wolves (Canis lupus) and arctic foxes (Alopex lagopus) are the only canid species found throughout the mainland tundra and arctic islands of North America. Contrasting evolutionary histories, and the contemporary ecology of each species, have combined to produce their divergent population genetic characteristics. Arctic foxes are more variable than wolves, and both island and mainland fox populations possess similarly high microsatellite variation. These differences result from larger effective population sizes in arctic foxes, and the fact that, unlike wolves, foxes were not isolated in discrete refugia during the Pleistocene. Despite the large physical distances and distinct ecotypes represented, a single, panmictic population of arctic foxes was found which spans the Svalbard Archipelago and the North American range of the species. This pattern likely reflects both the absence of historical population bottlenecks and current, high levels of gene flow following frequent long-distance foraging movements. In contrast, genetic structure in wolves correlates strongly to transitions in habitat type, and is probably determined by natal habitat-biased dispersal. Nonrandom dispersal may be cued by relative levels of vegetation cover between tundra and forest habitats, but especially by wolf prey specialization on ungulate species of familiar type and behaviour (sedentary or migratory). Results presented here suggest that, through its influence on sea ice, vegetation, prey dynamics and distribution, continued arctic climate change may have effects as dramatic as those of the Pleistocene on the genetic structure of arctic canid species.

The arctic fox (Alopex lagopus) in Scandinavia is classified as critically endangered after having gone through a severe decline in population size in the beginning of the 20th century, from which ...

Comparative analysis of the biology of insular and mainland populations has demonstrated a number of differences in characteristics of insular populations, termed the ‘island syndrome’. A subspecies of Arctic fox on the Commander Islands (Alopex lagopus semenovi on Mednyi Island) has been isolated for an evolutionarily significant time in small territories at the periphery of the species' range. The number of foxes on Mednyi had been observed to be very high since the islands were discovered in 1741, but a drastic decline in population density in the late 1970s, owing to mange, has left the population low. The aim of the study was to determine whether the Mednyi Arctic fox population exhibited the features expected in an isolated insular population, such as difference in body size, increased population density, larger social groups, lower tendency to disperse and lower fertility, and whether any behavioural changes were evolutionarily reinforced or were a temporal response to current ecological factors on the island. Eight predictions were identified based on the island syndrome, of which the Mednyi Arctic foxes conformed to seven. We suggest a new prediction, namely that the tendency for increased sex dimorphism in dispersal may also be a distinguishing feature of insular populations. All the features displayed by the insular population on Mednyi – conservative use of space, increased tendency to form complex groups, decreased fertility and dispersal – seemed to be preserved regardless of the currently comparatively low populations. Thus, although foxes have been below carrying capacity for at least 10 generations, island syndrome characteristics have persisted.

Swift foxes (Vulpes velox) are one of the smallest and most den-dependent and nocturnal canids in North America (Egoscue 1979, Tannerfeldt et al. 2003)Though research concerning swift fox fecundity has been conducted, it is a difficult and arduous task because pups do not emerge from natal dens until they are weaned at 5-7 weeks of age (Egoscue 1979). Previous techniques for determining fecundity of swift foxes have included excavating the den and counting pre-emergent pups (Cutter 1958, Kilgore 1969) and visual counts of post-emergent pups (Covell 1992, Olson and Lindzey 2002, Schauster et al. 2002, Harrison 2003, Sovada et al. 2003). Visual counts are the most commonly reported technique for counting post-emergent swift fox pups (Covell 1992, Olson and Lindzey 2002, Schauster et al. 2002, Harrison 2003, Sovada et al. 2003). In this technique >1 concealed observers with binoculars or spotting scopes watch a female's den from a distance (usually >40 m) in the evening and morning hours during early summer, in an attempt to count the number of pups. Visual counts also have been effectively used with similar species such as the kit fox (Vulpes macrotis; Cypher et al. 2000, Koopman et al. 2000) and arctic fox (V.lagopus; Strand et al. 2000). Prior research concerning determination of preemergent pups has been highly invasive, as dens were excavated or disturbed via observer presence. As more advanced technology becomes available to researchers, less insidious means of obtaining data can be implemented. Recently, video probe systems have been used to document various cavity-nesting and burrowing animals such as red-cockaded woodpeckers (Picoides borealis; Richardson et al. 1999) and western burrowing owls (Athene cunicularia hypugea; Gervais et al. 2000). Night-vision devices have been used to enumerate migrating owls (Tyto alba,Aegolius acadicus,Asio otus, Asio flammeus, and Otus asio; Russell et al. 1991) and to observe the emergence of big brown bats (Eptesicus fuscus) from maternal colonies (Kirkwood and Cartwright 1993). Automated videography has been used to observe nests and index population abundance (York 1991, Kristan et al. 1996, Booms and Fuller 2003). Yet, none of this technology has been evaluated for use in swift fox research. Measuring fecundity is an essential component in examining demographics of any wildlife species. Information on litter sizes is necessary to understand the dynamics of swift fox populations. Yet, no pups were observed from preliminary efforts at visual counts of swift fox pups during our study. Thus, the objective of this study was to compare visual counts, night vision, a den-probe system, and an automated video monitoring system for counting swift fox pups.

This study examines how the distribution and abundance of prey species influenced arctic fox Alopex lagopus diet on Svalbard during the summers of 1997, 1998 and 1999. The arctic fox together with the glaucous gull Larus hyperboreus are the only predators of this terrestrial ecosystem, and during summer the potential prey are confined to a few prey species such as various colonial seabirds, geese and Svalbard reindeer Rangifer tarandus platyrhynchus. There is great variation in distribution of prey over short distances, resulting in highly contrasting patterns in terms of both prey abundance and availability. Arctic fox diet was investigated by collecting scats (N = 818) at dens used for breeding. A cluster analysis based on similarities in the contents of scats revealed that arctic fox habitat could be divided into three distinct prey resource landscapes: two inland areas dominated by the prey species 'reindeer' and 'reindeer and geese', respectively, and one coastal resource area dominated by '...

This study investigated the physiological adaptations to fasting using the farmed blue fox (Alopex lagopus) as a model for the endangered wild arctic fox. Sixteen blue foxes were fed throughout the winter and 32 blue foxes were fasted for 22 d in Nov–Dec 2002. Half of the fasted blue foxes were food-deprived again for 22 d in Jan–Feb 2003. The farmed blue fox lost weight at a slower rate (0.97–1.02% body mass d−1) than observed previously in the arctic fox, possibly due to its higher initial body fat content. The animals experienced occasional fasting-induced hypoglycaemia, but their locomotor activity was not affected. The plasma triacylglycerol and glycerol concentrations were elevated during phase II of fasting indicating stimulated lipolysis, probably induced by the high growth hormone concentrations. The total cholesterol, HDL- and LDL-cholesterol, urea, uric acid and total protein levels and the urea:creatinine ratio decreased during fasting. Although the plasma levels of some essential amino acids increased, the blue foxes did not enter phase III of starvation characterized by stimulated proteolysis during either of the 22-d fasting procedures. Instead of excessive protein catabolism, it is liver dysfunction, indicated by the increased plasma bilirubin levels and alkaline phosphatase, alanine aminotransferase and aspartate aminotransferase activities, that may limit the duration of fasting in the species. J. Exp. Zool. 305A:32–46, 2006. © 2005 Wiley-Liss, Inc.

The aim of the study was to investigate the circannual rhythms of leptin and ghrelin in the blue fox, a variant of the endangered arctic fox, in relation to its seasonal cycles of body mass, adiposity and food intake. The effects of long-term fasting and exogenous melatonin treatment on these weight-regulatory hormones were also investigated. The leptin concentrations of the blue fox increased during the autumnal accumulation of fat and decreased during the wintertime and vernal weight loss periods. The leptin levels peaked 2-6 weeks before the maximum values were observed for the body mass indices, voluntary food intake, and body masses. The ghrelin concentrations fluctuated widely during the autumn but decreased in the winter in association with suppression of food intake. Exogenous melatonin advanced the seasonal changes in the food intake of the blue fox but did not affect the seasonal rhythms of leptin and ghrelin concentrations. The leptin concentrations did not respond to the 3-week fasting periods in a consistent way, but the ghrelin levels increased due to food deprivation. In addition to the amount of fat in the body the leptin secretion of the blue fox may be regulated also by other factors. The blue fox may also express seasonal changes in its leptin sensitivity. Our results reinforce the hypothesis that leptin does not function as an acute indicator of body adiposity in seasonal carnivores but rather as a long-term signal of nutritional status.

Summary 1. Home range size, spatial organization and territoriality of reproductive Arctic foxes were studied during the summer. The influence of spatial distribution and availability of the main prey was investigated in order to evaluate whether the spatial organization of Arctic foxes was coherent with key predictions of the resource dispersion hypotheses (RDH). The RDH includes the spatial characteristics of resource abundance, while there is also growing attention to the importance of the temporal characteristics of resource abundance. Hence the role of temporal and spatio-temporal predictability of prey explaining carnivore spatial organization was also investigated in this study. 2. The study was conducted on Svalbard; a simple High-Arctic terrestrial ecosystem which allowed unique estimates of prey abundance. The main prey of the Arctic fox (Svalbard reindeer Rangifer tarandus platyrhynchus , seabirds Alcidae and Procellariidae and geese Anseridae ) was surveyed systematically. These surveys revealed highly contrasting patterns in prey abundance within the terrestrial ecosystem. 3. Arctic fox summer home ranges varied in size (4‐60 km 2 ), as well as in overlap (17‐ 76%). The diverse spatial organization covaried with spatial and temporal patterns in prey abundance. Small home ranges (10 ± 5·6 km 2 ) with large overlap (76 ± 19·6%) were characteristic for coastal areas where prey was concentrated in small patches and predictable both in space and time. Medium home ranges (23 ± 4·2 km 2 ) and overlap (50 ± 6·6%) occurred inland where prey was clumped in larger patches and less predictable. Large home ranges (52 ± 8·4 km 2 ) with little overlap (17 ± 3·5%) occurred inland where prey was widely scattered and unpredictable. 4. Spatial dispersion and richness of prey resources explained most of the variation in Arctic fox spatial organization. The RDH framework could be used to explain the presence of relaxed territoriality found in this study. We suggest that the observed absence of more permanent social groups is due to the extremely severe winter conditions which force juvenile individuals to disperse from the natal area during the first winter. 5. Predictability of resources was an additional significant factor affecting both home-range size and overlap. Resource predictability captures the degree to which an animal can depend on its environment to offer suitable and secure living conditions over time. 6. This study emphasize the need to incorporate both spatial and temporal characteristics of resource distribution in order to fully understand the diversity of spatial arrangements among carnivores.

We determined the validity of water content, rump fat thickness, a subjective fat index, a kidney fat index, and an index based on total body weight (TBW) and the hind foot length (HFL) in order to predict lipid content of arctic foxes Alopex lagopus. The total fat content was either determined by extracting lipid from minced carcasses (N = 75) or by dissecting all visible adipose tissue (N = 35). Water content and rump fat thickness proved to be the best predictors of fat content in arctic foxes. The ratio TBW/HFL was also a reliable index and should be a convenient and practicable way of estimating the condition of live foxes. A subjective fat index could be applied when divided into three categories, whereas the kidney fat index was not a reliable predictor of lipid content of arctic foxes.

Summary 1. Terrestrial predators in coastal areas are often subsidized by marine foods. In order to determine the potential impact on terrestrial prey, the numerical response of predators to each food source must be determined. 2. In winter, arctic foxes ( Alopex lagopus ) may forage on the frozen Arctic ocean and scavenge carcasses of seals killed by polar bears ( Ursus maritimus ), but the importance of this food source and its effect on the population cycles of arctic foxes and lemmings (their primary prey) are unclear. 3. I estimated the marine component of the late winter diet of arctic foxes near Churchill, Manitoba, using stable-carbon isotope analysis, and compared these estimates to abundance of arctic foxes and collared lemmings ( Dicrostonyx richardsoni ). 4. From 1994 to 1997, fox density varied with lemming abundance, but following a decline, fox abundance began increasing before lemmings. During this increase marine foods were consumed more than in other years, with over two-thirds of food intake from marine sources. 5. Arctic and red fox ( Vulpes vulpes ) harvests in the 1980s to 1990s were correlated with published estimates of polar bear body mass, which varies with seal productivity. However, this relationship disappeared during high lemming years. 6. Thus, variation in marine productivity affects arctic fox abundance, especially when their primary prey are scarce, and this numerical response of arctic foxes to marine resources and lemmings suggests that increased predation by arctic foxes subsidized by seal carrion may delay the recovery of low lemming populations.

Summary This study compared polymorphism of nine canine-derived microsatellites (MS) (CPH1, CPH3, CPH6, CPH11, 2004, 2010, 2140, 2168 and 2319) in three species of the family Canidae. The DNA samples of 151 dogs, 53 arctic foxes and 91 red foxes were examined. The canine-derived primers did not amplify two MS (CPH1 and CPH11) in genome samples of the arctic fox. The most polymorphic MS in the studied species was the one named 2319. For majority of the loci mean allele size was higher in the dog than in two fox species. The genetic distances between the species were estimated using three formulas: DS, Da and (σμ)2. The estimated distances between both fox species were the smallest, while those between the dog and the arctic fox were the largest.

Seventy-seven arctic fox dens in Vindelfjallen, Northern Sweden, are described with regard to distribution, morphology and fox use. The density of dens in the area was 1 den / 21 km2 and dens were ...

A combined molecular and phenotypic approach was used to determine whether ear mites of the genus Otodectes (Acari: Psoroptidae) belong to a single species. The second internal transcribed spacer (ITS 2) of the rDNA of 16 isolates from 11 cats, two dogs, one arctic fox and two ferrets originating from four different continents was characterized. In addition, mites from dog, cat and arctic fox were investigated morphologically. Sequence comparisons revealed five different, but closely related genotypes which did not segregate according to host species or geographical origin. Morphologically, mites of the three host species did not differ significantly in their body or leg sizes. These investigations support the view that ear mites of the genus Otodectes from different hosts and geographical origins belong to a single species, Otodectes cynotis (Hering).

The arctic island group of Svalbard has no native species of voles or lemmings, but the vole Microtus rossiaemeridionalis has been accidentally introduced. Mainly found in the region of the abandoned Russian mining town of Grumant, the vole has not been able to colonise larger parts of Svalbard. The food habits of arctic foxes Alopex lagopus were studied by collecting and analysing samples of scats from various sites in the region in 1996, including fresh scats from three litters. Hairs of voles were found in scats from all sites, but the proportion of voles in the diet of arctic foxes varied considerably. Overall, voles made up 13% of the diet by occurrence, but were insignificant in the diet of the three litters. Birds were the most important (79%) and reindeer carcasses the second most important food item (22%). Other estimates used gave less importance to voles (10 and 4%). Birds most often constituted the major part of a scat, voles less often. Pups from three litters had consumed 97% birds, with the proportion of alcids and gulls varying largely according to availability near the respective litter. The population of voles at Grumant was very low in 1996, but in other years it may grow to much higher numbers. However, a large proportion of the scats collected were old and reflected arctic fox diet over many years. Although arctic foxes at Grumant have gained access to a new food source, voles appear to be of minor importance in their diet.

Mammals occupying northern areas may experience reduced fitness through the deleterious effects of snow on locomotion, energy expenditure, and food acquisition. Accordingly, in areas of deep and soft snow, selection may favour mammals possessing disproportionally large feet and lower footload (body mass/foot surface area). We collected carcasses of coyote Canis latrans (n = 472), red fox Vulpes vulpes (n = 199), arctic fox Alopex lagopus (n = 52), grey fox Urocyon cinereoargenteus (n = 17), and wolf Canis lupus (n = 14) and compared body mass, total foot area, difference in surface area between front and hind feet, and footload, among species and populations. All measured morphological attributes differed significantly among canid species; grey fox had the smallest feet, whereas arctic fox had both the lightest mass and lowest footload of all canids. For all species, adult males tended to be heavier and have larger feet than did adult females, and for most species adults were heavier and larger than were juveniles. Foot area in red foxes increased significantly with latitude, with populations found north of 488N (i.e. approximate latitude where snow may become limiting) averaging 12% larger feet than those found farther south. For coyotes, body mass increased with latitude, with populations found north of 488N averaging 26% heavier mass than those occurring farther south. Coyote foot area also increased with latitude, with populations found north of 488N averaging 25% larger feet than those found farther south. When indices of snow severity (mean annual snowfall, mean number of days with snowcover) were considered, foot area for red foxes was correlated with annual snowfall as well as latitude, whereas coyote morphology failed to correlate with any snow-related variables. These results suggest that snow may have contributed to selection for foot size in some wild canids (i.e. arctic fox, red fox), but such selective forces were probably relatively weak and inconsistent across species.

Many hypotheses have been raised to try explaining the failure of Fennoscandian arctic fox Alopex lagopus populations to recover following 60–70 years of protection. One of the most appealing hypotheses has been that they depend on scavenging the kills of ungulates made by large carnivores, mainly wolves Canis lupus, which have been absent for much of the 20th century. We examine this hypothesis using a variety of data from throughout the holarctic range of arctic foxes and conclude that there is little evidence for the importance of this food source - although it is an issue that needs to be examined in greater detail.

Reliable and abundant resources are likely to favour specialization, while unpredictable environmental variation should favour a generalist strategy. The rodent population cycles of northern latitudes can be seen as both predictable and unpredictable, depending on the scale in time and space. The arctic fox Alopex lagopus is an opportunistic carnivore, but paradoxically, it seems to function as a specialist on fluctuating rodent Arvicolinae populations in most inland areas. We have studied the dietary response of arctic foxes in Sweden during 5 years of varying abundance of Norwegian lemming Lemmus lemmus, and how these changes influenced the reproductive success of the foxes. The arctic fox population on mainland Fennoscandia is threatened by extinction and the situation has deteriorated during the 1980s and 1990s because of an absence of lemming peaks. Our results showed that in all years, lemming was the main prey for arctic foxes, with 85% frequency of occurrence in summer faeces (scats). Bird remains (mainly Passeriformes) were present in 34% of the scats, reindeer Rangifer tarandus in 21%, voles and shrews in 4% and hares Lepus timidus in 2% of the scats. The occurrences of lemming, bird and larger mammal (reindeer and hare) remains in the scats varied significantly between years. Temporal variations within summer seasons and dietary differences between sub-areas, indicated that arctic foxes fed opportunistically on the alternative prey types. Den occupancy rates were positively correlated with lemming population densities during the previous winter, indicating a strong numerical response. We conclude that from a functional aspect, the arctic fox in Sweden is a lemming specialist, since lemming is the main prey and their abundance is the best predictor of arctic fox reproductive success. Other prey are used opportunistically in relation to their availability.