Adaptation by the Arctic Fox (<i>Alopex lagopus</i>) to the Polar Winter

Metabolic rates of four resting, post-absorptive male adult summer- and winter-adapted captive arctic foxes (Alopex lagopus) were recorded. Basal metabolic rates (BMR) varied seasonally with a 36% increase from winter to summer, while body mass was reduced by 17% in the same period. The lower critical temperature (T 1c) of the winter-adapted arctic fox was estimated to −7°C, whereas T lc during summer was 5°C. The similarity of these values, which are much higher than hitherto assumed (e.g. Scholander et al. 1950b), is mainly due to a significantly (P<0.05) lower BMR in winter than in summer. Body core (stomach) temperature was stable, even at ambient temperatures as low as −45°C, but showed a significant (P<0.05) seasonal variation, being lower in winter (39.3±0.33°C) than in summer (39.8±0.16°C). The thermal conductivity of arctic fox fur was the same during both seasons, whereas the thermal conductance in winter was lower than in summer. This was reflected in an increase in fur thickness of 140% from summer to winter, and in a reduced metabolic response to ambient temperatures below T lc in winter. Another four arctic foxes were exposed to three periods of forced starvation, each lasting 8 days during winter, when body mass is in decline. No significant reduction in mass specific BMR was observed during the exposure to starvation, and respiratory quotient was unchanged at 0.73±0.02 during the first 5 days, but dropped significantly (P<0.05) to 0.69±0.03 at day 7. Locomotor activity and body core (intraperitoneal) temperature was unaltered throughout the starvation period, but body mass was reduced by 18.5±2.1% during these periods. Upon re-feeding, locomotor activity was significantly (P<0.05) reduced for about 6 days. Energy intake was almost doubled, but stabilised at normal levels after 11 days. Body mass increased, but not to the level before the starvation episodes. Instead, body mass increased until it reached the reduced body mass of ad libitum fed control animals. This indicates that body mass in the arctic fox is regulated according to a seasonally changing set point.

Responses of one species to climate change may influence the population dynamics of others, particularly in the Arctic where food webs are strongly linked. Specifically, changes to the cryosphere may limit prey availability for predators. We examined Arctic (Vulpes lagopus) and red fox (V. vulpes) population dynamics near the southern edge of the Arctic fox distribution using fur harvest records from Churchill, Manitoba, Canada between 1955 and 2012. Arctic foxes showed a declining population trend over time (inferred from harvest records corrected for trapping effort), whereas the red fox population trend was relatively stable. The positive relationship between the annual Arctic and red fox harvests suggested interspecific competition did not promote the Arctic fox decline. To investigate alternative mechanisms, we evaluated the relative influence of sea-ice phenology, snow depth, snow duration, winter thaws, and summer temperature on the harvest dynamics of both species in the most recent 32years (1980-2012; n = 29) of our data. Arctic fox harvests were negatively related to the length of time Hudson Bay was free of sea ice. Shorter sea ice duration may reduce access to seal carrion as an alternative winter food source when lemming densities decline. Contrary to our prediction, red fox harvest was not related to summer temperature but was positively related to snow depth, suggesting winter prey availability may limit red fox population growth. Predators have an important ecological role, so understanding the influence of changes in the cryosphere on predator-prey interactions may better illuminate the broader influence of climate change on food-web dynamics.

In the Arctic tundra, predators face recurrent periods of food scarcity and often turn to ungulate carcasses as an alternative food source. As important and localized resource patches, carrion promotes co-occurrence of different individuals, and its use by predators is likely to be affected by interspecific competition. We studied how interspecific competition and resource availability impact winter use of carrion by Arctic and red foxes in low Arctic Fennoscandia. We predicted that the presence of red foxes limits Arctic foxes' use of carrion, and that competition depends on the availability of other resources. We monitored Arctic and red fox presence at supp lied carrion using camera traps. From 2006 to 2021, between 16 and 20 cameras were active for 2 months in late winter (288 camera-winters). Using a multi-species dynamic occupancy model at a week-to-week scale, we evaluated the use of carrion by foxes while accounting for the presence of competitors, rodent availability, and supplemental feeding provided to Arctic foxes. Competition affected carrion use by increasing both species' probability to leave occupied carcasses between consecutive weeks. This increase was similar for the two species, suggesting symmetrical avoidance. Increased rodent abundance was associated with a higher probability of colonizing carrion sites for both species. For Arctic foxes, however, this increase was only observed at carcasses unoccupied by red foxes, showing greater avoidance when alternative preys are available. Supplementary feeding increased Arctic foxes' carrion use, regardless of red fox presence. Contrary to expectations, we did not find strong signs of asymmetric competition for carrion in winter, which suggests that interactions for resources at a short time scale are not necessarily aligned with interactions at the scale of the population. In addition, we found that competition for carcasses depends on the availability of other resources, suggesting that interactions between predators depend on the ecological context.

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 (21years). 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.

Metabolic rate and evaporative water loss at different ambient temperatures in two species of fox: The red fox ( Vulpes vulpes) and the arctic fox ( Alopex lagopus)

... The main objectives of my work are to examine (1) how arctic foxes use seasonally abundant foods and (2) how seasonal and annual fluctuations in food abundance affect foraging behaviours and population dynamics of arctic foxes. I am especially interested in how arctic foxes use geese and their eggs (i.e., seasonally abundant foods) and how this varies with fluctuations in small mammal abundance (i.e., foods that fluctuate annually). ... My work is done at Karrak Lake (67°14'N, 100°16'W) and surrounding areas in the Queen Maud Gulf Bird Sanctuary in Nunavut, Canada. ... Fieldwork for my project was done in the spring and summers of 2000-04, and data analyses are currently underway. I monitor population dynamics of arctic foxes in two goose nesting areas at Karrak Lake and two areas outside the influence of nesting geese, whereas I monitor foraging behaviours of arctic foxes in one section of the goose colony at Karrak Lake. ... I examine foraging behaviours of arctic foxes by observing individually marked foxes with spotting scopes .... Avoiding cache loss to competitors is a critical component for the evolution of caching .... I examine how nesting distribution of geese and dispersal of geese away from the colony affect cache loss by evaluating the survival rate of experimentally deployed caches .... I examine arctic fox diets by comparing isotope ratios (delta 13C and delta 15N) of fox tissues ... with those of food items collected in the field .... Fur is metabolically inactive, whereas blood is metabolized continuously ..., so by examining spring blood and winter fur I obtain information on both spring and fall diets. Geese are not present at Karrak Lake in either spring or fall, so goose signatures in this study represent foods cached in summer. I monitor population dynamics of arctic foxes ... through line-transect surveys, mark-recapture studies, and den inventories. ... Goose eggs (from both nests and existing caches) made up 91% of all foods taken by arctic foxes during goose-nesting at Karrak Lake. Foxes cached 96% of these eggs (i.e., most eggs from existing caches were moved to new locations) whereas other foods (i.e., small mammals, geese, and passerine eggs) were either consumed immediately or brought back to den sites. ... Age of cache sites and dispersal by geese away from the colony after hatch (when ca. 1 million geese and their offspring leave the colony in about 10 days) affected loss of experimental caches. These results suggest that both food abundance and strategies to prevent ageing of cache sites (e.g., cache site selection and moving of caches in poor condition) were important in affecting the arrangement of caches at our study site. Cached eggs constituted 30% to 40% of the arctic foxes' diet in autumn and 0% to 30% in spring. ... The abundance of arctic foxes was predominantly affected by abundance of geese ... whereas the density of breeding foxes and litter size were predominantly affected by small mammal abundance. ... This study will provide information on how seasonal and annual fluctuations in food abundance influence use of stored foods and population dynamics of arctic foxes. ... This study will also provide information on predator-prey interactions, which can be used for management and conservation of both arctic foxes and Arctic-nesting birds. ...

A large number of analyses have examined how basal metabolic rate (BMR) is affected by body mass in mammals. By contrast, the critical ambient temperatures that define the thermo-neutral zone (TNZ), in which BMR is measured, have received much less attention. We provide the first phylogenetic analyses on scaling of lower and upper critical temperatures and the breadth of the TNZ in 204 mammal species from diverse orders. The phylogenetic signal of thermal variables was strong for all variables analysed. Most allometric relationships between thermal variables and body mass were significant and regressions using phylogenetic analyses fitted the data better than conventional regressions. Allometric exponents for all mammals were 0.19 for the lower critical temperature (expressed as body temperature - lower critical temperature), -0.027 for the upper critical temperature, and 0.17 for the breadth of TNZ. The small exponents for the breadth of the TNZ compared to the large exponents for BMR suggest that BMR per se affects the influence of body mass on TNZ only marginally. However, the breadth of the TNZ is also related to the apparent thermal conductance and it is therefore possible that BMR at different body masses is a function of both the heat exchange in the TNZ and that encountered below and above the TNZ to permit effective homeothermic thermoregulation.

Snow cover has dramatic effects on the structure and functioning of Arctic ecosystems in winter. In the tundra, the subnivean space is the primary habitat of wintering small mammals and may be critical for their survival and reproduction. We have investigated the effects of snow cover and habitat features on the distributions of collared lemming (Dicrostonyx groenlandicus) and brown lemming (Lemmus trimucronatus) winter nests, as well as on their probabilities of reproduction and predation by stoats (Mustela erminea) and arctic foxes (Vulpes lagopus). We sampled 193 lemming winter nests and measured habitat features at all of these nests and at random sites at two spatial scales. We also monitored overwinter ground temperature at a subsample of nest and random sites. Our results demonstrate that nests were primarily located in areas with high micro-topography heterogeneity, steep slopes, deep snow cover providing thermal protection (reduced daily temperature fluctuations) and a high abundance of mosses. The probability of reproduction increased in collared lemming nests at low elevation and in brown lemming nests with high availability of some graminoid species. The probability of predation by stoats was density dependent and was higher in nests used by collared lemmings. Snow cover did not affect the probability of predation of lemming nests by stoats, but deep snow cover limited predation attempts by arctic foxes. We conclude that snow cover plays a key role in the spatial structure of wintering lemming populations and potentially in their population dynamics in the Arctic.

Based on studies (2016-2022) of diphyllobothriasis infestation of animals in the territory of the Commander Islands National Park, it was revealed that the average extent of Bering Arctic Fox invasion was 37.3%, with seasonal fluctuations: winter – 42.7%, spring – 36.6%, summer – 34.9% (fecal helminthoscopy, n=351). The American mink is infected with diphyllobothriids with an invasion intensity of 26.5%. Helminthological autopsies (n=11) revealed strobils of Diphyllobothrium spp. - 5-15 specimens, 20-70 cm long, in 5 Bering Arctic foxes. At the autopsy of 10 individuals of the American mink and 1 individual of the Mednovsky Arctic fox, no diphyllobothriids were found, as well as during helminthoscopic examination of the feces of the Mednovsky Arctic fox (n=37). Encapsulated plerocercoids were detected in the spawning individuals of Oncorhynchus gorbusha from the territory of Bering Island. Key words: National Park "Commander Islands", diphyllobothriids, prevalence, Bering Arctic fox, Mednovsky Arctic fox, American mink.

cultured fibroblasts of cell lines from animals of different size have similar activities for a range of different enzymes and metabolic rates that converge on the similar values despite the metabolic rates of the animals of different sizes having quite distinctly different basal metabolic rates (

Selection for crypsis has been recognized as an important ecological driver of animal colouration, whereas the relative importance of thermoregulation is more contentious with mixed empirical support. A potential thermal advantage of darker individuals has been observed in a wide range of animal species. Arctic animals that exhibit colour polymorphisms and undergo seasonal colour moults are interesting study subjects for testing the two alternative hypotheses: demographic performance of different colour morphs might be differentially affected by snow cover with a cryptic advantage for lighter morphs, or conversely by winter temperature with a thermal advantage for darker morphs. In this study, we explored whether camouflage and thermoregulation might explain differences in reproduction and survival between the white and blue colour morphs of the Arctic fox Vulpes lagopus under natural conditions. Juvenile and adult survival, breeding propensity and litter size were measured for 798 captive-bred and released or wild-born Arctic foxes monitored during an 11-year period (2007-2017) in two subpopulations in south-central Norway. We investigated the proportion of the two colour morphs and compared their demographic performance in relation to spatial variation in duration of snow cover, onset of snow season and winter temperatures. After population re-establishment, a higher proportion of blue individuals was observed among wild-born Arctic foxes compared to the proportion of blue foxes released from the captive population. Our field study provides the first evidence for an effect of colour morph on the reproductive performance of Arctic foxes under natural conditions, with a higher breeding propensity of the blue morph compared to the white one. Performance of the two colour morphs was not differentially affected by the climatic variables, except for juvenile survival. Blue morph juveniles showed a tendency for higher survival under colder winter temperatures but lower survival under warmer temperatures compared to white morph juveniles. Overall, our findings do not consistently support predictions of the camouflage or the thermoregulation hypotheses. The higher success of blue foxes suggests an advantage of the dark morph not directly related to disruptive selection by crypsis or thermoregulation. Our results rather point to physiological adaptations and behavioural traits not necessarily connected to thermoregulation, such as stress response, immune function, sexual behaviour and aggressiveness. Our findings highlight the need to explore the potential role of genetic linkage or pleiotropy in influencing the fitness of white and blue Arctic foxes as well as other species with colour polymorphisms.

This work was conducted to determine effect of season and starvation on metabolic rate during running in the Arctic Fox (Alopex lagopus) on Svalbard (78°55’N, 11°56’E), Norway. Indirect calorimetry was used to measure metabolic rate of foxes running on a treadmill and heart rate was monitored using implanted radio transmitters. The relationship between heart rate and metabolic rate was also examined. Metabolic rate increased with running speed. In July the metabolic rate during running almost fitted general equations predicted for mammals, while it was up to 20% lower in January, indicating seasonal variation in metabolic rate. There was a significant positive linear relationship between heart rate and weight specific metabolic rate, suggesting that heart rate can be used as an indicator of metabolic rate. Starvation for 11 days decreased the net cost of running by 13% in January and 17% in July, suggesting that a starved fox runs more energetically efficient than when fed. Heart rate measured in July decreased by 27% during starvation. Re-feeding reversed the starvation-induced reduction in metabolic rate and heart rate during running almost up to post-absorptive levels. The present results are from one fox, and must be considered as preliminary data until further studies are conducted.

Post-absorptive resting metabolic rates (RMRs), body mass and ad libitum food intake were recorded on an annual cycle in captive arctic foxes (Alopex lagopus) at Svalbard. During the light season in May and in the dark period in November, RMR during starvation and subsequent re-feeding were also measured. In contrast to earlier findings, the present study indicated a seasonal trend in post-absorptive RMR (in W.kg-1 and W.kg-0.75). The values in the light summer were 15% and 11% higher than the values in the dark winter, suggesting a physiological adaptation aiding energy conservation during winter in arctic foxes. Body mass and ad libitum food intake varied inversely through the year. A significant reduction in RMR (in W and W.kg-0.75) with starvation (metabolic depression) was recorded both in May and November, indicating an adaptation to starvation in arctic foxes. The lack of metabolic depression during a period of starvation that was concomitant with extremely cold ambient temperatures in November 1994 indicates that metabolic responses to starvation may be masked by thermoregulatory needs. At very low ambient temperatures, arctic foxes may require increased heat production which cannot be achieved via below-average rates of metabolism.

Food-caching by arctic foxes ( Vulpes lagopus (L., 1758)) is a behavioural adaptation thought to increase winter survival, especially in bird colonies where a large number of eggs can be cached during a short nesting season. In this paper, we measured the energy content of greater snow goose ( Chen caerulescens atlantica Kennard, 1927) eggs and evaluated their perishability when cached in tundra soil for a whole summer. We estimated that eggs lost only ~8% of their dry mass over 60 days of storage in the ground. We used published estimates on digestibility of nutrients by arctic foxes to estimate that fresh and stored goose eggs contained 816 and 730 kJ of metabolizable energy, respectively, a difference of 11%. Using information on arctic fox energetics, we evaluated that 145 stored eggs were required to sustain the growth of one pup from the age of 1 to 3 months (nutritional independence). Moreover, 23 stored eggs were energetically equivalent to the average fat deposit of an arctic fox during winter. Finally, we calculated that an adult arctic fox would need to recover 160–220 stored eggs to survive 6 months in resting conditions during cold winter temperatures. This value increased to 480 when considering activity cost. Based on egg acquisition and caching rates observed in many goose colonies, we conclude that cached eggs represent an important source of energy relative to the needs of an arctic fox during winter, and have thus a high fitness value.

Physical characteristics of 73 arctic fox (Alopex lagopus) dens in Svalbard are described. In a mountainous study area of 975 sq/km, most dens were found below 200 m and none was found above 400 m. Most dens were located in slopes in valleys or along the coast, facing in a southerly direction (mean aspect 222 degrees). Dens had an average of 9.8 entrances and a mean size of 52.1 sq/m. Snow cover was less over dens than on adjacent areas in winter. Vegetation at dens was different than at adjacent areas. Of 56 dens found in the study area, 3 were burrows and the rest were situated under boulders, in screes or in bedrock. Dens were situated in dry localities, most often in protruding bedrock and ridges where foxes had an unrestricted view.Key words: arctic fox, Alopex lagopus, dens, Norway, Svalbard