Natal den selection by sympatric arctic and red foxes on Herschel Island,Yukon, Canada

In the twentieth century, red fox (Vulpes vulpes) expanded into the Canadian Arctic, where it competes with arctic fox (Vulpes lagopus) for food and shelter. Red fox dominates in physical interactions with the smaller arctic fox, but little is known about competition between them on the tundra. On Hershel Island, north Yukon, where these foxes are sympatric, we focused on natal den choice, a critical aspect of habitat selection. We tested the hypothesis that red fox displaces arctic fox from dens in prey-rich habitats. We applied an approach based on model comparisons to analyse a 10-year data set and identify factors important to den selection. Red fox selected dens in habitats that were more prey-rich in spring. When red foxes reproduced, arctic fox selected dens with good springtime access, notably many burrows unblocked by ice and snow. These provided the best refuge early in the reproductive season. In the absence of red foxes, arctic foxes selected dens offering good shelter (i.e. large isolated dens). Proximity to prey-rich habitats was consistently less important than the physical aspects of dens for arctic fox. Our study shows for the first time that red foxes in the tundra select dens associated primarily with prey-rich areas, while sympatric arctic foxes do not. These results fit a model of red fox competitively interfering with arctic fox, the first detailed study of such competition in a true arctic setting.     

Exclusion by interference competition? The relationship between red and arctic foxes

The distribution of many predators may be limited by interactions with larger predator species. The arctic fox in mainland Europe is endangered, while the red fox is increasing its range in the north. It has been suggested that the southern distribution limit of the arctic fox is determined by interspecific competition with the red fox. This has been criticised, on the basis that the species co-exist on a regional scale. However, if the larger red fox is superior and interspecific competition important, the arctic fox should avoid close contact, especially during the breeding season. Consequently, the distribution of breeding dens for the two species would be segregated on a much smaller spatial and temporal scale, in areas where they are sympatric. We tested this hypothesis by analysing den use of reproducing arctic and red foxes over 9 years in Sweden. High quality dens were inhabited by reproducing arctic foxes more often when no red foxes bred in the vicinity. Furthermore, in two out of three cases when arctic foxes did reproduce near red foxes, juveniles were killed by red foxes. We also found that breeding arctic foxes occupied dens at higher altitudes than red foxes did. In a large-scale field experiment, red foxes were removed, but the results were not conclusive. However, we conclude that on the scale of individual territories, arctic foxes avoid areas with red foxes. Through interspecific interference competition, the red fox might thus be excluding the arctic fox from breeding in low altitude habitat, which is most important in years when food abundance is limited and competition is most fierce. With high altitude refuges being less suitable, even small-scale behavioural effects could scale up to significant effects at the population level.     

Rabies and canine distemper in an arctic fox population in Alaska

Two oil field workers were attacked by a rabid arctic fox (Alopex lagopus) in the Prudhoe Bay oil field (Alaska, USA) prompting officials to reduce the local fox population. Ninety-nine foxes were killed during winter 1994. We tested foxes for prevalence of rabies and canine distemper. Exposure to rabies was detected in five of 99 foxes. Of the five, only one fox had rabies virus in neural tissue as determined by the direct fluorescent antibody test. The other four foxes had been exposed to rabies, but had apparently produced antibodies and did not have an active infection. No evidence of canine distemper was detected as determined by the absence of distemper antibodies in serum and distemper virus in neural tissue.     

Genetic polymorphism of plasma α1B-glycoprotein and transferrin in arctic and silver foxes

Plasma samples of 235 foxes from 38 complete families (14 of arctic foxes, 21 of silver foxes and 3 with arctic x silver fox hybrid offspring) were analysed by one-dimensional horizontal polyacrylamide gel electrophoresis (PAGE) pH 9.0 followed by general-protein staining of gels. A major postalbumin of fox plasma was identified as alpha 1B-glycoprotein (alpha 1B) by using immunoblotting with antiser m specific to human or pig plasma alpha 1B. Four codominant, autosomal alleles of alpha 1B were found in arctic foxes. Two transferrin (TF) alleles (TfF, TfS) were observed in arctic foxes and two (TfD, Tff) in silver foxes; the TF F type of both of the fox species showed identical electrophoretic mobilities. The arctic foxes showed a high degree of polymorphism for both TF and alpha 1B. The silver foxes showed a scarce polymorphism of TF and were monomorphic for alpha 1B. The arctic fox, silver fox and their hybrids could be clearly differentiated from one another by their plasma protein patterns obtained by the PAGE method.     

The breeding productivity of dark-bellied brent geese and curlew sandpipers in relation to changes in the numbers of arctic foxes and lemmings on the Taimyr Peninsula, Siberia —

There were about three-year cycles in the populations of arctic foxes, and the breeding productivities of brent geese and curlew sandpipers on the Taimyr Peninsula, Russia, The populations of arctic foxes and lemmings changed in synchrony. The breeding productivities of the birds tended to be good when the arctic foxes were increasing in numbers and poor when the arctic foxes were decreasing. There was a negative relationship between arctic fox numbers (or occupied lairs) and the breeding productivity of brent geese in the following year. Although there was evidence of wide-spread synchrony In the lemming cycle across the Taimyr Peninsula, some localities showed differences, However, such sites would still have been influenced by the general pattern of fox abundance in the typical tundra zone of the Taimyr Peninsula, where most of the arctic foxes breed and from which extensive movements of foxes occur after a decline in lemming numbers. The results support a prey-switching hypothesis (also known as the alternative prey hypothesis) whereby arctic foxes, and other predators, feed largely on lemmings when these are abundant or increasing, but switch to birds when the lemming population is small or declining. The relationships between arctic foxes, lemmings and brent geese may be further influenced by snowny owls which create fox-exclusion zones around their nests, thus providing safe nesting areas for the geese.     

Historical and ecological determinants of genetic structure in arctic canids

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.     

Population structure of two rabies hosts relative to the known distribution of rabies virus variants in Alaska

For pathogens that infect multiple species, the distinction between reservoir hosts and spillover hosts is often difficult. In Alaska, three variants of the arctic rabies virus exist with distinct spatial distributions. We tested the hypothesis that rabies virus variant distribution corresponds to the population structure of the primary rabies hosts in Alaska, arctic foxes (Vulpes lagopus) and red foxes (Vulpes vulpes) to possibly distinguish reservoir and spillover hosts. We used mitochondrial DNA (mtDNA) sequence and nine microsatellites to assess population structure in those two species. mtDNA structure did not correspond to rabies virus variant structure in either species. Microsatellite analyses gave varying results. Bayesian clustering found two groups of arctic foxes in the coastal tundra region, but for red foxes it identified tundra and boreal types. Spatial Bayesian clustering and spatial principal components analysis identified 3 and 4 groups of arctic foxes, respectively, closely matching the distribution of rabies virus variants in the state. Red foxes, conversely, showed eight clusters comprising two regions (boreal and tundra) with much admixture. These results run contrary to previous beliefs that arctic fox show no fine‐scale spatial population structure. While we cannot rule out that the red fox is part of the maintenance host community for rabies in Alaska, the distribution of virus variants appears to be driven primarily by the arctic fox. Therefore, we show that host population genetics can be utilized to distinguish between maintenance and spillover hosts when used in conjunction with other approaches.     

Prevalence and geographical distribution of the ear canker mite (Otodectes cynotis) among arctic foxes (Alopex lagopus) in Iceland

Three hundred forty five adult arctic foxes (Alopex lagopus) from all counties in Iceland were examined for excess cerumen and ear canker mites (Otodectes cynotis). Only 13 foxes (4%) from a single county in northwestern Iceland were infested, where the prevalence of otodectiasis was 38%. Whether or not this parasite is new to the arctic fox in Iceland is unknown. If it is recently introduced, possible sources of infestation are farmed silver foxes (Vulpes vulpes), domestic dogs, domestic or feral cats, and arctic foxes from Greenland. It appears that the rate of transmission between adult foxes is low; a more common route of transmission is probably from the mother to her offspring or between vixens breeding in the same dens in subsequent years by contamination of the dens. No correlation was found between the prevalence of mites in foxes and Samson character.     

Northern nomads: ability for extensive movements in adult arctic foxes

In July 2008 we outfitted reproductively active adult arctic foxes with satellite tracking collars on Bylot Island, Nunavut, Canada and recorded their movements over a complete annual cycle. We present the tracking data from two individuals, one female and one male, who traveled extensively from February to July 2009, covering minimum distances of 4,599 and 2,193 km, respectively. We recorded high and sustained travel rates on both land and sea ice that reached 90 km/day for the female and 88 km/day for the male. Our data confirm that arctic foxes can move extensively and demonstrate sustained travel rates that are 1.5 times those previously measured for the species. Our study is the first presenting detailed year-round satellite tracking of adult arctic foxes and has implications for our understanding of navigational abilities, foraging ecology, trophic interactions with lemming populations, and genetic population structure of arctic foxes.     

Prevalence of Toxoplasma gondii infection diagnosed by PCR in farmed red foxes, arctic foxes and raccoon dogs

The aim of this study was to compare Toxoplasma gondii infection in three canid species: red fox Vulpes vulpes, arctic fox Vulpes lagopus and raccoon dog Nyctereutesprocyonoides kept at the same farm. Anal swabs were taken from 24 adult and 10 juvenile red foxes, 12 adult arctic foxes, three adult and seven juvenile raccoon dogs. Additionally, muscle samples were taken from 10 juvenile red foxes. PCR was used to detect T. gondii DNA. T. gondii infection was not detected in any of the arctic foxes; 60% ofraccoon dogs were infected; the prevalence of the parasite in material from red fox swabs was intermediate between the prevalence observed in arctic foxes and raccoon dogs. It is possible that susceptibility and immune response to the parasite differ between the three investigated canid species. T. gondii DNA was detected in muscle tissue of five young foxes. The results of this study suggest that T. gondii infection is not rare in farmed canids.     

Detection of farm fox and hybrid genotypes among wild arctic foxes in Scandinavia

In Scandinavia, farmed arctic foxes frequently escape from farms, raising concern about hybridization with the endangered wild population. This study was performed to find a genetic marker to distinguish escaped farm foxes from wild Scandinavian foxes. Microsatellite and mitochondrial control region variation were analyzed in 41 farm foxes. The results were compared with mitochondrial and microsatellite data from the wild population in Scandinavia. The farm foxes were genetically distinct from the wild foxes (F _ST=0.254, P < 0.00001) and all farm foxes had a single control region haplotype different from those observed in the wild population. We developed a method based on Restriction Fragment Length Polymorphism (RFLP) on the mitochondrial control region to differentiate between farmed and wild arctic foxes. This test was subsequently successfully used on 25 samples from free-ranging foxes, of which four had a suspected farm origin. All four of the suspected foxes, and none of the others, carried the farm fox haplotype. Three of these were successfully genotyped for all eleven microsatellite loci. A population assignment test and a Bayesian Markov Chain Monte Carlo analysis indicated that two of these individuals were escaped farm foxes, and that the third possibly was a hybrid between a farmed and a wild arctic fox.     

Farmed arctic foxes on the Fennoscandian mountain tundra: implications for conservation

Hybridization between wild and captive-bred individuals is a serious conservation issue that requires measures to prevent negative effects. Such measures are, however, often considered controversial by the public, especially when concerning charismatic species. One of the threats to the critically endangered Fennoscandian arctic fox Alopex lagopus is hybridization with escaped farm foxes, conveying a risk of outbreeding depression through loss of local adaptations to the lemming cycle. In this study, we investigate the existence of escaped farm foxes among wild arctic foxes and whether hybridization has occurred in the wild. We analysed mitochondrial control region sequences and 10 microsatellite loci in samples from free-ranging foxes and compared them with reference samples of known farm foxes and true Fennoscandian arctic foxes. We identified the farm fox specific mitochondrial haplotype H9 in 25 out of 182 samples, 21 of which had been collected within or nearby the wild subpopulation on Hardangervidda in south-western Norway. Genetic analyses of museum specimens collected on Hardangervidda (1897–1975) suggested that farm fox genotypes have recently been introduced to the area. Principal component analysis as well as both model- and frequency-based analyses of microsatellite data imply that the free-ranging H9s were farm foxes rather than wild arctic foxes and that the entire Hardangervidda population consisted of farm foxes or putative hybrids. We strongly recommend removal of farm foxes and hybrids in the wild to prevent genetic pollution of the remaining wild subpopulations of threatened arctic foxes.     

Experimental Encephalitozoonosis in the Blue Fox

Newborn and young pups up to the age of 15 days were exposed to E. cuniculi, either by keeping the pups in cages together with orally inoculated foster-mothers and their offspring, or by oral inoculation with E. cuniculi spores. A majority of pups appeared sero-positive to E. cuniculi with the india-ink immuno-reaction from 35 to 87 days post exposure; spores of E. cuniculi were detected in organs of some of the animals. The non-inoculated pups kept together with the orally inoculated pups became seropositive from 49 to 129 days after the oral inoculations. However, the exposure of newborn and young pups failed to induce clinical encephalitozoo-nosis, and when killed at the time of pelting the body weights and fur quality appeared to be within the normal range in all exposed foxes. No macroscopic lesions were detected in the various organs. Histologically focal interstitial nephritis occurred in the great majority of the seropositive animals. Meningoencephalitis was seen in some of the foxes, whereas slightly thickened walls of some arteries, mainly in the myocardium, were found in a few animals. The lesions of the brain and kidneys seem to be very similar to those seen in chronic cases of rabbit encephalitozoonosis. Polyarteritis nodosa and severe encephalitis and interstitial nephritis with extensive proliferations of plasma cells, which are almost constant findings in cases of clinically diseased foxes, were not detected in any of the subclinically infected animals. Various factors that might be of significance in the pathogenesis of the disease are discussed, and it is concluded that intrauterine infection of the pups via the transplacental route appears to be an essential supposition for the establishment of clinical fox encephalitozoonosis.     

Age-dependent genetic structure of arctic foxes in Svalbard

Arctic foxes are highly mobile arctic predators with a very weak population genetic structure over large parts of their range. Less is, however, known about the more local genetic structure within regions. Here, we analyze genotypes at 12 microsatellite loci for 561 arctic foxes trapped in the high-arctic archipelago Svalbard and investigate the genetic structure in three different age classes. Significant linkage disequilibrium, deficit of heterozygotes, genetic differentiation, and a decrease in relatedness with distance among animals trapped in their first winter suggested that some litter mates remain in proximity of each other during the first winter. This pattern was stronger for females than for males, indicating male-biased juvenile dispersal, and weaker for older animals. There was no genetic differentiation among adult foxes harvested in different hunting areas. The foxes from the protected area around Hornsund were however more differentiated than expected based on geographic distance alone, suggesting a possible disrupting effect of harvest on the spatial genetic structure in the rest of Svalbard. Our results also indicated a possible kin structure among adult females, suggesting natal philopatry, but further investigations will be needed to reach firm conclusions concerning kin structure in arctic foxes.     

Sea-ice use by arctic foxes in northern Alaska

The extensive use of sea-ice by three arctic foxes (Alopex lagopus) in northern Alaska was documented using satellite telemetry during the winter of 2005–2006. Here we present the first detailed data on movements of individual foxes while on the sea-ice. Two juvenile males and one juvenile female traveled long distances (904, 1,096, and 2,757 km) and remained on the sea-ice for extended periods of time (76, 120, and 156 days). Average distances traveled per day ranged from 7.5 to 17.6 km and foxes achieved maximum rates of travel of up to 61 km/day. These findings verify the use of sea-ice by arctic foxes and raise concerns that the diminishing arctic ice cover may negatively impact populations by limiting access to marine food sources.     

DNA analysis on fox faeces and competition induced niche shifts

Interference competition can force inferior competitors to change their distribution patterns. It is, however, possible that the dominant competitor poses a higher threat during certain times of the year, for example during reproduction. In such cases, the inferior competitor is expected to change its distribution accordingly. We used a molecular species identification method on faeces to investigate how the spatial overlap between arctic and red foxes changes between seasons. The results show that arctic and red foxes are sympatric during winter, but allopatric in summer as arctic foxes retreat to higher altitudes further from the tree-line during the breeding season.     

Predator–prey relationships: arctic foxes and lemmings

1. The number of breeding dens and litter sizes of arctic foxes Alopex lagopus were recorded and the diet of the foxes was analysed during a ship-based expedition to 17 sites along the Siberian north coast. At the same time the cyclic dynamics of co-existing lemming species were examined.
2. The diet of arctic foxes was dominated by the Siberian lemming Lemmus sibiricus (on one site the Norwegian lemming L. lemmus ), followed by the collared lemming Dicrostonyx torquatus .
3. The examined Lemmus sibiricus populations were in different phases of the lemming cycle as determined by age profiles and population densities.
4. The numerical response of arctic foxes to varying densities of Lemmus had a time lag of 1 year, producing a pattern of limit cycles in lemming–arctic fox interactions. Arctic fox litter sizes showed no time lag, but a linear relation to Lemmus densities. We found no evidence for a numerical response to population density changes in Dicrostonyx .
5. The functional or dietary response of arctic foxes followed a type II curve for Lemmus , but a type III response curve for Dicrostonyx .
6. Arctic foxes act as resident specialist for Lemmus and may increase the amplitude and period of their population cycles. For Dicrostonyx , on the other hand, arctic foxes act as generalists which suggests a capacity to dampen oscillations.     

The impact of past climate change on genetic variation and population connectivity in the Icelandic arctic fox

Previous studies have suggested that the presence of sea ice is an important factor in facilitating migration and determining the degree of genetic isolation among contemporary arctic fox populations. Because the extent of sea ice is dependent upon global temperatures, periods of significant cooling would have had a major impact on fox population connectivity and genetic variation. We tested this hypothesis by extracting and sequencing mitochondrial control region sequences from 17 arctic foxes excavated from two late-ninth-century to twelfth-century AD archaeological sites in northeast Iceland, both of which predate the Little Ice Age (approx. sixteenth to nineteenth century). Despite the fact that five haplotypes have been observed in modern Icelandic foxes, a single haplotype was shared among all of the ancient individuals. Results from simulations within an approximate Bayesian computation framework suggest that the rapid increase in Icelandic arctic fox haplotype diversity can only be explained by sea-ice-mediated fox immigration facilitated by the Little Ice Age.     

Use of Supplementary Feeding Dispensers by Arctic Foxes in Norway

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. © 2020 The Authors. Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.