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Species that seasonally moult from brown to white to match snowy backgrounds become conspicuous and experience increased predation risk as snow cover duration declines. Long-term adaptation to camouflage mismatch in a changing climate might occur through phenotypic plasticity in colour moult phenology and or evolutionary shifts in moult rate or timing. Also, adaptation may include evolutionary shifts towards winter brown phenotypes that forgo the winter white moult. Most studies of these processes have occurred in winter white populations, with little attention to polymorphic populations with sympatric winter brown and winter white morphs. Here, we used remote camera traps to record moult phenology and mismatch in two polymorphic populations of Arctic foxes in Sweden over 2 years. We found that the colder, more northern population moulted earlier in the autumn and later in the spring. Next, foxes moulted earlier in the autumn and later in the spring during colder and snowier years. Finally, white foxes experienced relatively low camouflage mismatch while blue foxes were mismatched against snowy backgrounds most of the autumn through the spring. Because the brown-on-white mismatch imposes no evident costs, we predict that as snow duration decreases, increasing blue morph frequencies might help facilitate species persistence.

Understanding and predicting large-scale ecological responses to global environmental change requires comparative studies across geographic scales with coordinated efforts and standardized methodologies. We designed, applied and assessed standardized protocols to measure tundra herbivory at three spatial scales: plot, site (habitat), and study area (landscape). The plot and site-level protocols were tested in the field during summers 2014-2015 at eleven sites, nine of them comprising warming experimental plots included in the International Tundra Experiment (ITEX). The study area protocols were assessed during 2014-2018 at 24 study areas across the Arctic. Our protocols provide comparable and easy-to-implement methods for assessing the intensity of invertebrate herbivory within ITEX plots and for characterizing vertebrate herbivore communities at larger spatial scales. We discuss methodological constraints and make recommendations for how these protocols can be used and how sampling effort can be optimized to obtain comparable estimates of herbivory, both at ITEX sites and at large landscape scales. The application of these protocols across the tundra biome will allow characterizing and comparing herbivore communities across tundra sites and at ecologically relevant spatial scales, providing an important step towards a better understanding of tundra ecosystem responses to large-scale environmental change. CGB was funded by the Estonian Research Council (grant IUT 20-28), and the European Regional Development Fund (Centre of Excellence EcolChange). JDMS was supported by the Research Council of Norway (262064). OG and LB were supported by the French Polar Institute (program “1036 Interactions”) and PRC CNRS Russie 396 (program “ICCVAT”). DSH, NL, MAG, JB and JDR were supported by the Natural Sciences and Engineering Research Council (Canada). NL, MAG, JB and JDR were supported by the Polar Continental Shelf Program. NL was supported by the Canada Research Chair program and the Canada Foundation for Innovation. NL and JB were supported by Environment Canada and Polar Knowledge Canada. NL and MAG were supported by the Government of Nunavut, the Igloolik Community, and Université de Moncton. NL, MAG and JB were supported by the Northern Scientific Training Program. JMA was funded by Carl Tryggers stiftelse för vetenskaplig forskning and Qatar Petroleum (QUEX-CAS-QP-RD-18_19). IHM-S was funded by the UK Natural Environmental Research Council Shrub Tundra (NE/M016323/1) grant. ISJ was funded by the University of Iceland Research Fund. Fieldwork in Yamal peninsula (Erkuta, Sabetta and Belyi) for DE, NS and AS was supported by the Russian Foundation for Basic Research (No: 18-05-60261 and No: 18-54-15013), Fram Centre project YaES (No: 362259), the Russian Center of Development of the Arctic, and the “Yamal-LNG” company. Fieldwork in Utqiaġvik was supported by the U.S. Fish and Wildlife Service. Fieldwork in Svalbard was supported by the Norwegian Research Council (AFG No: 246080/E10), the Norwegian Polar Institute, Climate-ecological Observatory for Arctic Tundra – COAT, the Svalbard Environmental protection fund (project number 15/20), and the University Centre in Svalbard (UNIS) and the AB-338/AB-838 students of 2018. Sampling at Billefjorden was supported by GACR 17- 20839S.

The grey seal ( Halichoerus grypus) has been part of the Baltic Sea fauna for more than 9000 years and has ever since been subjected to extensive human hunting, particularly during the early phases of its presence in the Baltic Sea, but also in the early 20th century. In order to study their temporal genetic structure and to investigate whether there has been a genetically continuous grey seal population in the Baltic, we generated mitochondrial control region data from skeletal remains from ancient grey seals from the archaeological sites Stora Förvar (Sweden) and Neustadt (Germany) and compared these with modern grey seal data. We found that the majority of the Mesolithic grey seals represent haplotypes that is not found in contemporary grey seals, indicating that the Baltic Sea population went extinct, likely due to human overexploitation and environmental change. We hypothesize that grey seals recolonised the Baltic Sea from the North Sea. during the Bronze Age or Iron Age, and that the contemporary Baltic grey seal population is direct descendants of this recolonisation. Our study highlights the power of biomolecular archaeology to understand the factors that shape contemporary marine diversity.

The hypothesis that predation is the cause of the regular small rodent population oscillations observed in boreal and Arctic regions has long been debated. Within this hypothesis, it is proposed that the most likely predators to cause these destabilizing effects are sedentary specialists, with small mustelids being possible candidates. One such case would be the highly specialized least weasel (Mustela nivalis) driving the Norwegian lemming (Lemmus lemmus) cycle in Fennoscandia. These predators are often elusive and therefore distribution data can only be based on field signs, which is problematic when various mustelid species are sympatric, such as weasels and stoats (Mustela erminea). Here we present the results of using mustelid faeces in predated winter lemming nests to correctly identify the predator and thus discern which species exerts the strongest predation pressure on lemming winter populations. Samples were obtained during different phases in the lemming cycle, spanning 6 years, to account for different prey densities. Faecal mitochondrial DNA extraction and amplification of a 400-bp fragment was successful in 92/114 samples (81%); the sequencing of these samples proved that most predation occurrences (83%) could be attributed to the least weasel. These findings support the hypothesis that weasels in particular show high specificity in predation and could therefore be candidates to driving the lemming cycle in this area. We conclude that DNA analysis of faecal remains around predated nests can be a useful tool for further investigations concerning predator–prey interactions in the tundra.

Accelerating climate change is causing severe habitat fragmentation in the Arctic, threatening the persistence of many cold-adapted species. The Scandinavian arctic fox (Vulpes lagopus) is highly fragmented, with a once continuous, circumpolar distribution, it struggled to recover from a demographic bottleneck in the late 19th century. The future persistence of the entire Scandinavian population is highly dependent on the northernmost Fennoscandian subpopulations (Scandinavia and the Kola Peninsula), to provide a link to the viable Siberian population. By analyzing 43 arctic fox genomes, we quantified genomic variation and inbreeding in these populations. Signatures of genome erosion increased from Siberia to northern Sweden indicating a stepping-stone model of connectivity. In northern Fennoscandia, runs of homozygosity (ROH) were on average ~1.47-fold longer than ROH found in Siberia, stretching almost entire scaffolds. Moreover, consistent with recent inbreeding, northern Fennoscandia harbored more homozygous deleterious mutations, whereas Siberia had more in heterozygous state. This study underlines the value of documenting genome erosion following population fragmentation to identify areas requiring conservation priority. With the increasing fragmentation and isolation of Arctic habitats due to global warming, understanding the genomic and demographic consequences is vital for maintaining evolutionary potential and preventing local extinctions.

Accelerating climate change is causing severe habitat fragmentation in the Arctic, threatening the persistence of many cold-adapted species. The Scandinavian arctic fox (

Reduced fitness through genetic drift and inbreeding is a major threat to small and isolated populations. Although previous studies have generally used genetically verified pedigrees to document effects of inbreeding and gene flow, these often fail to capture the whole inbreeding history of the species. By assembling a draft arctic fox (Vulpes lagopus) genome and resequencing complete genomes of 23 additional foxes born before and after a well-documented immigration event in Scandinavia, we here look into the genomic consequences of inbreeding and genetic rescue. We found a difference in genome-wide diversity, with 18% higher heterozygosity and 81% lower FROH in immigrant F1 compared to native individuals. However, more distant descendants of immigrants (F2, F3) did not show the same pattern. We also found that foxes with lower inbreeding had higher probability to survive their first year of life. Our results demonstrate the important link between genetic variation and fitness as well as the transient nature of genetic rescue. Moreover, our results have implications in conservation biology as they demonstrate that inbreeding depression can effectively be detected in the wild by a genomic approach.

The Arctic fox (Vulpes lagopus L.) population on the Kola Peninsula occupies an intermediate, and potentially connecting, position between foxes living on the Scandinavian Peninsula and populations further east in Russia, but very little is known about the status of this population. Here we summarize data from the literature, forgotten archival sources about research in the first half of the twentieth century, and the results of several independent expeditions undertaken over the past two decades. These materials include data on fur harvesting, incomplete monitoring data from official winter track counts of game animals, local knowledge, and our own observations. Our research revealed the extremely poor state of the Arctic fox population on the Kola Peninsula. According to our estimates, the current population is likely isolated and consists of no more than a few dozen adults. The fur return data, together with long-term data on small rodent abundance, suggest that irregular and fading out lemming cycles were a major driver of the Arctic fox population decline. The thorough research from the 1930s contrasts strongly with the lack of interest in studying and monitoring the population in recent decades, which is not even listed as a threatened species in the regional Red Data Book. In fact, the work performed here filled a more than a half-century gap in the study of the population and allowed us to determine the urgent need to resume research and immediately take active measures to protect and promote the recovery of the species in the region.

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 ...

In socially flexible species, the tendency to live in groups is expected to vary through a trade-off between costs and benefits, determined by ecological conditions. The Resource Dispersion Hypothesis predicts that group size changes in response to patterns in resource availability. An additional dimension is described in Hersteinsson’s model positing that sociality is further affected by a cost–benefit trade-off related to predation pressure. In the arctic fox (Vulpes lagopus), group-living follows a regional trade-off in resources’ availability and intra-guild predation pressure. However, the effect of local fluctuations is poorly known, but offers an unusual opportunity to test predictions that differ between the two hypotheses in systems where prey availability is linked to intra-guild predation. Based on 17-year monitoring of arctic fox and cyclic rodent prey populations, we addressed the Resource Dispersion Hypothesis and discuss the results in relation to the impact of predation in Hersteinsson’s model. Group-living increased with prey density, from 7.7% (low density) to 28% (high density). However, it remained high (44%) despite a rodent crash and this could be explained by increased benefits from cooperative defence against prey switching by top predators. We conclude that both resource abundance and predation pressure are factors underpinning the formation of social groups in fluctuating ecosystems.

Most birds incubate their eggs to allow embryo development. This behaviour limits the ability of adults to perform other activities. Hence, incubating adults trade-off incubation and nest protection with foraging to meet their own needs. Parents can either cooperate to sustain this trade-off or incubate alone. The main cause of reproductive failure at this reproductive stage is predation and adults reduce this risk by keeping the nest location secret. Arctic sandpipers are interesting biological models to investigate parental care evolution as they may use several parental care strategies. The three main incubation strategies include both parents sharing incubation duties (?biparental?), one parent incubating alone (?uniparental?), or a flexible strategy with both uniparental and biparental incubation within a population (?mixed?). By monitoring the incubation behaviour in 714 nests of seven sandpiper species across 12 arctic sites, we studied the relationship between incubation strategy and nest predation. First, we described how the frequency of incubation recesses (NR), their mean duration (MDR), and the daily total duration of recesses (TDR) vary among strategies. Then, we examined how the relationship between the daily predation rate and these components of incubation behaviour varies across strategies using two complementary survival analysis. For uniparental and biparental species, the daily predation rate increased with the daily total duration of recesses and with the mean duration of recesses. In contrast, daily predation rate increased with the daily number of recesses for biparental species only. These patterns may be attributed to two independent mechanisms: cryptic incubating adults are more difficult to locate than unattended nests and adults departing the nest or feeding close to the nest can draw predators' attention. Our results demonstrate that incubation behaviour as mediated by incubation strategy has important consequences for sandpipers? reproductive success. This article is protected by copyright. All rights reserved.

Lemmings are a key component of tundra food webs and changes in their dynamics can affect the whole ecosystem. We present a comprehensive overview of lemming monitoring and research activities, and assess recent trends in lemming abundance across the circumpolar Arctic. Since 2000, lemmings have been monitored at 49 sites of which 38 are still active. The sites were not evenly distributed with notably Russia and high Arctic Canada underrepresented. Abundance was monitored at all sites, but methods and levels of precision varied greatly. Other important attributes such as health, genetic diversity and potential drivers of population change, were often not monitored. There was no evidence that lemming populations were decreasing in general, although a negative trend was detected for low arctic populations sympatric with voles. To keep the pace of arctic change, we recommend maintaining long-term programmes while harmonizing methods, improving spatial coverage and integrating an ecosystem perspective. ELECTRONIC SUPPLEMENTARY MATERIAL: The online version of this article (10.1007/s13280-019-01198-7) contains supplementary material, which is available to authorized users.

Human view on nature has fluctuated over time, depending on contemporary knowledge and beliefs. In recent centuries, the view has shifted from an instrumental to an existential apprehension of natu ...

Waders that breed in the sub-Arctic are one of the groups most threatened by climate change. At the same time, wader breeding success also can vary as a function of fluctuations in the numbers of predators and rodents (an alternative prey for the predators). How climate change could influence these foodweb interactions remains poorly studied. In this study, we analysed the effects of ecological (e.g. vole/lemming and predator abundance) and environmental factors (e.g. snow cover) on the breeding success of waders in sub-Arctic Lapland. We monitored more than 500 wader nests during six breeding seasons, which spanned a full rodent cycle and one year of exceptionally late snow melt. Nest predation rate, and thus wader breeding success, did not vary as a function of predator or rodent abundance. However, predation rate was exceptionally high in the year with a late snow melt. More variability in climate is expected for the future, where more precipitation and cold spring temperatures resulting in late snow melt will be more frequent, influencing the rodent and predator numbers, and therefore wader breeding success in the sub-Arctic. Snow would limit the number of open areas for nesting and hence predators would then be able to find these nests more easily. Additionally, predators might concentrate their efforts on alternative prey if snow has reduced their capacity to find other food sources. And, ultimately, changes in the rodent fluctuations could affect the final outcome of predators.

This is a post-peer-review, pre-copyedit version of an article published in Ambio. The final authenticated version is available online at: http://dx.doi.org/10.1007/s13280-019-01198-7. Lemmings are a key component of tundra food webs and changes in their dynamics can affect the whole ecosystem. We present a comprehensive overview of lemming monitoring and research activities, and assess recent trends in lemming abundance across the circumpolar Arctic. Since 2000, lemmings have been monitored at 49 sites of which 38 are still active. The sites were not evenly distributed with notably Russia and high Arctic Canada underrepresented. Abundance was monitored at all sites, but methods and levels of precision varied greatly. Other important attributes such as health, genetic diversity and potential drivers of population change, were often not monitored. There was no evidence that lemming populations were decreasing in general, although a negative trend was detected for low arctic populations sympatric with voles. To keep the pace of arctic change, we recommend maintaining long-term programmes while harmonizing methods, improving spatial coverage and integrating an ecosystem perspective.

Most birds incubate their eggs, which requires time and energy at the expense of other activities. Birds generally have two incubation strategies: biparental where both mates cooperate in incubating eggs, and uniparental where a single parent incubates. In harsh and unpredictable environments, incubation is challenging due to high energetic demands and variable resource availability. We studied the relationships between the incubation behaviour of sandpipers (genus Calidris) and two environmental variables: temperature and a proxy of primary productivity (i.e. NDVI). We investigated how these relationships vary between incubation strategies and across species among strategies. We also studied how the relationship between current temperature and incubation behaviour varies with previous day's temperature. We monitored the incubation behaviour of nine sandpiper species using thermologgers at 15 arctic sites between 2016 and 2019. We also used thermologgers to record the ground surface temperature at conspecific nest sites and extracted NDVI values from a remote sensing product. We found no relationship between either environmental variables and biparental incubation behaviour. Conversely, as ground-surface temperature increased, uniparental species decreased total duration of recesses (TDR) and mean duration of recesses (MDR), but increased number of recesses (NR). Moreover, small species showed stronger relationships with ground-surface temperature than large species. When all uniparental species were combined, an increase in NDVI was correlated with higher mean duration, total duration and number of recesses, but relationships varied widely across species. Finally, some uniparental species showed a lag effect with a higher nest attentiveness after a warm day while more recesses occurred after a cold day than was predicted based on current temperatures. We demonstrate the complex interplay between shorebird incubation strategies, incubation behaviour, and environmental conditions. Understanding how species respond to changes in their environment during incubation helps predict their future reproductive success.

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

Genome-wide association studies provide good opportunities for studying the genetic basis of adaptive traits in wild populations. Yet, previous studies often failed to identify major effect genes. In this study, we used high-density single nucleotide polymorphism and individual fitness data from a wild non-model species. Using a whole-genome approach, we identified the MC1R gene as the sole causal gene underlying Arctic fox Vulpes lagopus fur colour. Further, we showed the adaptive importance of fur colour genotypes through measures of fitness that link ecological and evolutionary processes. We found a tendency for blue foxes that are heterozygous at the fur colour locus to have higher fitness than homozygous white foxes. The effect of genotype on fitness was independent of winter duration but varied with prey availability, with the strongest effect in years of increasing rodent populations. MC1R is located in a genomic region with high gene density, and we discuss the potential for indirect selection through linkage and pleiotropy. Our study shows that whole-genome analyses can be successfully applied to wild species and identify major effect genes underlying adaptive traits. Furthermore, we show how this approach can be used to identify knowledge gaps in our understanding of interactions between ecology and evolution.

Genetic rescue can facilitate the recovery of small and isolated populations suffering from inbreeding depression. Long-term effects are however complex, and examples spanning over multiple generations under natural conditions are scarce. The aim of this study was to test for long-term effects of natural genetic rescue in a small population of Scandinavian Arctic foxes (Vulpes lagopus). By combining a genetically verified pedigree covering almost 20 years with a long-term dataset on individual fitness (n = 837 individuals), we found no evidence for elevated fitness in immigrant F2 and F3 compared to native inbred foxes. Population inbreeding levels showed a fluctuating increasing trend and emergence of inbreeding within immigrant lineages shortly after immigration. Between 0–5 and 6–9 years post immigration, the average number of breeding adults decreased by almost 22% and the average proportion of immigrant ancestry rose from 14% to 27%. Y chromosome analysis revealed that 2 out of 3 native male lineages were lost from the gene pool, but all founders represented at the time of immigration were still contributing to the population at the end of the study period through female descendants. The results highlight the complexity of genetic rescue and suggest that beneficial effects can be brief. Continuous gene flow may be needed for small and threatened populations to recover and persist in a longer time perspective.

Understanding ecological relationships between humans and marine predators is crucial for the implementation of sustainable management practices. Comprehensive estimation of pinniped diet is essential for assessing interaction with fisheries and often has an important conservational value. Due to uncertainty regarding the accuracy of methods traditionally used to estimate harbour seal (Phoca vitulina) diet it is necessary to improve analysis methods. We investigated the diet of harbour seals hauling out in an estuary area in north-western Iceland between May and August of 2010 and 2011 by genetic (molecular) analysis of prey in faeces using DNA metabarcoding. The results were compared to previously published results from morphological analysis. Our results showed that species consumed were mainly sandeels (Ammodytes sp.), flatfishes (Pleuronectidae), gadoids (Gadidae), herring (Clupea harengus) and capelin (Mallotus villosus). The results from molecular and morphological analyses were similar in regards to important prey species, but species diversity was lower in the morphological analysis and 38% of the samples included prey items that were unidentifiable in the morphological analysis. Notably, despite Atlantic salmon (Salmo salar), brown trout (Salmo trutta) and Arctic char (Salvelinus alpinus) availability in the study area, neither of the methods found evidence of salmonids in the harbour seal diet. Recently, a severe decline has been observed in the Icelandic harbour seal population. Since the main reason for culling harbour seals in Iceland is to reduce predation on salmonids, findings presented in this paper have essential conservation implications and suggest that culling needs to be reassessed.

The grey wolf (Canis lupus Linnaeus, 1758) is one of the most widespread large carnivores on Earth, and occurs throughout the Arctic. Although wolf diet is well studied, we have scant information from high Arctic areas. Global warming is expected to increase the importance of predation for ecosystem regulation in Arctic environments. To improve our ability to manage Arctic ecosystems under environmental change, we therefore need knowledge about Arctic predator diets. Prey remains in 54 wolf scats collected at three sites in the high Arctic region surrounding the Hall Basin (Judge Daly Promontory, Ellesmere Island, Canada, and Washington Land and Hall Land, both in northwestern Greenland) pointed to a dietary importance of arctic hare (Lepus arcticus Ross, 1819; 55% frequency of occurrence) and muskoxen (Ovibos moschatus (Zimmermann, 1780); 39% frequency of occurrence), although we observed diet variation among the sites. A literature compilation suggested that arctic wolves (Canis lupus arctos Pocock, 1935) preferentially feed on caribou (Rangifer tarandus (Linnaeus, 1758)) and muskoxen, but can sustain themselves on arctic hares and Greenland collared lemmings (Dicrostonyx groenlandicus (Traill, 1823)) in areas with limited or no ungulate populations. We suggest that climate change may alter the dynamics among wolves, arctic hare, muskoxen, and caribou, and we encourage further studies evaluating how climate change influences predator–prey interactions in high Arctic environments.

Seal watching as a form of wildlife tourism is becoming increasingly popular worldwide. Behavioral changes caused by the presence of tourists could lead to negative consequences for seal welfare and may affect reproduction and survival. Therefore, managing seal-watching activities to ensure future protection and conservation is important. Codes of conduct or guidelines for how to behave around animals are one way to regulate wildlife watching and are often easier and quicker to implement than laws. Codes explaining the consequences for wildlife if the code is not followed appeal to the moral obligation of tourists and thereby increase incentives to act appropriately. This study focused on analyzing the content of codes of conduct for seal watching. Codes of conduct (n = 33) accessible on the internet during the time of study were analyzed. Results show that in many areas where seal watching occurs there are no regulations or guidelines. The content and detail of the codes varied and the information was often insufficient to offer adequate protection of seals. Few of the codes were developed in cooperation with scientists or stated that the content was based on research. Further, a majority of the codes did not explain the consequences for wildlife if the code was not followed. More research on seals and the tourists watching them is needed to better understand the effects of tourism and how disturbance could be minimized. Meanwhile, developing an international code of conduct (with local additions) built on existing knowledge in the field could be one option to increase protection and ensure conservation of these animals. The results presented in this article could assist the development of such a code of conduct.

Mating patterns are highly context-dependent and the outcome of selection pressures formed by ecological factors, inbreeding levels and access to available partners. In small and inbred populations, matings are limited by high kin encounter rates and access to mates. In this paper, we use background pedigree data to investigate mating patterns and inbreeding avoidance in an isolated and critically endangered Arctic fox (Vulpes lagopus) population. Empirical data showed avoidance of matings within natal family. Based on 35 documented matings, we only recorded two full-sibling matings and these occurred between individuals from different natal families. Matings between second-order relatives, however, occurred to the same extent as between unrelated individuals. To test how this influenced the population development of inbreeding (f), we simulated scenarios of random mating, exclusion of natal family and exclusion of individuals in already existing pair bonds. The observed development of inbreeding did not correspond the expected scenario of random mating (linear regression, r2 = 0.354, P = 0.20), but showed a comparable outcome as the simulated development of discriminating natal family (linear regression, r2 = 0.980, P

There are both positive and negative impacts on wildlife associated with wildlife tourism. In Sweden, the endangered Arctic fox is subject to a growing tourist interest. In the Helags mountain regi ...

The Swedish Polar Research Secretariat and National Science Foundation as part of the research program SWEDARCTIC 2016 to the project ‘Arctic Islands’ (2013-7812-106995-50). Further financial support was received from the Spanish Ministry of Economy and Competitiveness, the Natural Sciences and Engineering Research Council of Canada, the Canada Research Chair Program, the Canadian Foundation for Innovation, Northern and Arboriginal Affairs, Centre d’Etudes Nordiques, and the Swedish Research Council.

Reproductive experience affects juvenile survival in a wide range of species with possible links to differences in foraging capacity and predation. Using supplementary feeding, we aimed to limit direct effect of prey abundance to investigate indirect effects of small-rodent availability and maternal experience on juvenile summer survival rates in an endangered population of arctic fox (Vulpes lagopus (L., 1758)). We used data spanning 7 years, included a complete small-rodent cycle, comprising 49 litters and 394 cubs. The effect of small-rodent abundance on juvenile survival depended on maternal breeding experience. Cubs born by first-time-breeding females had lower survival rate when small-rodent abundance was low compared with juveniles born to experienced mothers who remained unaffected. It was unlikely due to starvation, as physical condition was unrelated to survival. Instead, we favour the explanation that intraguild predation was an important cause of mortality. There was a negative relationship between survival and amount of time cubs were left unattended, suggesting that parental behaviour affected predation. We propose that a prey switch related to small-rodent abundance caused fluctuations in intraguild predation pressure and that inexperienced females were less able to cope with predation when small rodents were scarce.

Run to the hills : gene flow among mountain areas leads to low genetic differentiation in the Norwegian lemming

The Arctic fox (Vulpes lagopus) population in Fennoscandia experienced a drastic bottleneck in the late 19th century as a result of high hunting pressure. In the 1990s, despite nearly 70 years of protection, the population showed no signs of recovery. In order to mitigate the population decline and facilitate re-establishment, conservation actions including supplementary feeding and red fox culling were implemented in 1998, followed by the reintroduction of foxes from a captive breeding programme, starting in 2006. A positive demographic impact of these actions is evident from a doubling of the population size over the past decade. We used genetic data collected in eight subpopulations between 2008 and 2015 to address whether the recent demographic recovery has been complemented by changes in genetic variation and connectivity between subpopulations. Our results show that genetic variation within subpopulations has increased considerably during the last decade, while genetic differentiation among subpopulations has decreased. A marked shift in metapopulation dynamics is evident during the study period, suggesting substantially increased migration across the metapopulation. This shift followed the recolonization of an extinct subpopulation through the release of foxes from the captive breeding programme and was synchronized in time with the implementation of supplementary feeding and red fox culling in stepping stone patches between core subpopulations in mid-Scandinavia. Indeed, the increase in genetic variation and connectivity in the Scandinavian arctic fox population suggests that metapopulation dynamics have been restored, which may indicate an increase in the long-term viability of the population. This is an open access article distributed under the terms of the Creative Commons CC-BY license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

Abstract Life history theory predicts that individuals will differ in their risk-taking behavior according to their expected future fitness. Understanding consequences of such individual variation within a behavioral trait is crucial in explaining potential trade-offs between different traits and in predicting future dynamics in changing environments. Here, we studied individuals in a wild arctic fox population to explore if (1) individual variation in risk-taking behaviors of adult arctic foxes and in stress-dealing behaviors of their juveniles exist and are consistent over time to verify the existence of personality traits; (2) those behavioral traits in adults and juveniles are correlated; (3) they can explain fitness-related components (i.e., juvenile physical condition, mortality rate). We presented simple field experiments assessing behavioral traits by observing adult reactions toward approaching observers, and juvenile behaviors while trapping. Through the experiments, we found highly consistent individual variation of adults in vigilance and boldness levels, and more flexible juvenile behavioral traits categorized as investigating, passive, and escaping. The offspring of bolder adults exhibited more investigating behaviors and were less passive than the offspring of shy adults. Juvenile physical condition was not related to their mortality nor any behavioral traits of either parents or themselves. Lastly, highly investigating and active juveniles with bold parents had significantly lower mortality rates. This shows that interactions between parent personality and juvenile behavioral traits affect a fitness-related component in the life history of individuals. Significance statement The recent surge of interest in consistent individual difference in behavior, also called as animal personality, has already focused on its fitness consequences, but few studies have investigated the interactions between parent and offspring personality, and their ecological consequences. Moreover, this has rarely been studied in wild canids. The arctic fox is a charismatic species showing wide individual variation in behaviors. They live in highly fluctuating tundra ecosystems providing different selection regimes, making it even more eco-evolutionarily intriguing. Yet, few studies looked into behavioral traits and their importance in this system. While introducing simple methods to improve personality research in the wild, we provide a unique example of how variation in both parents and their juveniles collectively works for group dynamics in a cyclic population. This provides a firm basic for understanding behavior-mediated dynamics and opens up broader questions on how fluctuating environments exert varying pressures on individual differences.

The rapidly growing interest for nature-based recreational activities threaten biodiversity values and increases the disturbance caused to wildlife. Several studies have demonstrated spatial and temporal activity shifts of animals in response to human disturbance. However, most studies investigate effects on how animals use an area of high tourism disturbance, such as a tourist resort or hiking trail, and not the effects of human disturbance at a key site for the animal, such as a denning or breeding site. In this study, we use photos from remote camera monitoring of arctic fox (Vulpes lagopus) dens with the aim to investigate the effects of various levels of tourism disturbance on the diurnal activity pattern and vigilance of breeding adult arctic foxes at the den site. We find a temporal shift towards a higher daytime activity at the den in response to high intensity tourism (71 ± 3.9% in disturbed areas compared with 53 ± 6.2% in undisturbed areas), which stands in contrast to an increased nocturnality seen in studies of many other species, including another study of arctic foxes. We suggest that the difference could be explained by the higher cost of avoiding a key site for the animal, as in this study, compared with avoiding a more general human disturbance in an area within the distribution range of the animal, as in most other studies. Increased time spent avoiding the perceived threat of humans could compromise other important activities and have potential negative effects on e.g. hunting or provisioning for the juveniles. Human disturbance focused at a key site, such as a denning or breeding site, can thereby be expected to have larger consequences than what is observed in most studies of disturbance effects on wildlife. Based on observations of both tourists and arctic foxes during close encounters in an area of high intensity tourism, we also find that they both respond behaviorally to each other. The potential for positive or negative feedback mechanisms in such relationships between tourists and wildlife highlights the importance of considering both sides of the complex interaction to find a balance between preserving biodiversity and ensuring continued possibilities for recreation.

Species distribution models (SDMs) are often calibrated using presence- only datasets plagued with environmental sampling bias, which leads to a decrease of model accuracy. In order to compensate for this bias, it has been suggested that background data (or pseudoabsences) should represent the area that has been sampled. However, spatially-explicit knowledge of sampling effort is rarely available. In multi-species studies, sampling effort has been inferred following the target-group (TG) approach, where aggregated occurrence of TG species informs the selection of background data. However, little is known about the species- specific response to this type of bias correction. The present study aims at evaluating the impacts of sampling bias and bias correction on SDM performance. To this end, we designed a realistic system of sampling bias and virtual species based on 92 terrestrial mammal species occurring in the Mediterranean basin. We manipulated presence and background data selection to calibrate four SDM types. Unbiased (unbiased presence data) and biased (biased presence data) SDMs were calibrated using randomly distributed background data. We used real and TG-estimated sampling efforts in background selection to correct for sampling bias in presence data. Overall, environmental sampling bias had a deleterious effect on SDM performance. In addition, bias correction improved model accuracy, and especially when based on spatially-explicit knowledge of sampling effort. However, our results highlight important species-specific variations in susceptibility to sampling bias, which were largely explained by range size: widely-distributed species were most vulnerable to sampling bias and bias correction was even detrimental for narrow-ranging species. Furthermore, spatial discrepancies in SDM predictions suggest that bias correction effectively replaces an underestimation bias with an overestimation bias, particularly in areas of low sampling intensity. Thus, our results call for a better estimation of sampling effort in multispecies system, and cautions the uninformed and automatic application of TG bias correction. This article is protected by copyright. All rights reserved.

Personality exists in non-human animals and can impact fitness. There is, however, a shortage of empirical studies in certain areas within the field, and fundamental evolutionary theory on personality remains largely untested. For example, little is known on how variation in personality is maintained over evolutionary time. Theory suggests that fluctuating selection pressures due to spatiotemporal variation in conditions, e.g. food availability, is a possible mechanism and a few studies have shown that the success of different personality types varies with spatiotemporal conditions. However, it remains unknown whether different mechanisms can maintain personality within a species. Here we use a reintroduction programme for the critically endangered European mink (Mustela lutreola) to test whether multiple personality trait domains (boldness, exploration and sociability) affected survival in two different years and islands. This was done through pre-release personality tests and post-release radio-tracking monitoring. Survival was positively correlated with boldness, whereas the relationship with exploration was either negative or positive depending on year/island. The results show a complex relationship between personality and survival and suggest that exploration can be maintained over evolutionary time via spatiotemporal variation in conditions. However, in contrast to exploration, boldness did not vary spatiotemporally and sociability had no impact on survival. This indicates that different personality trait domains might be maintained by different mechanisms. To date, personality has been studied primarily within behavioural sciences, but through empirical findings we highlight the importance of personality also in ecology and conservation biology. Electronic supplementary material The online version of this article (doi:10.1007/s00442-016-3740-0) contains supplementary material, which is available to authorized users.

Aim: The majority of the non-volant mammals now present on the island of Gotland, Sweden, have been introduced in modern times. One exception is the mountain hare (Lepus timidus), which was present ...

In territorial species, observed density dependence is often manifest in lowered reproductive output at high population density where individuals have fewer resources or are forced to inhabit low-quality territories. The Arctic fox (Vulpes lagopus) in Iceland is territorial throughout the year and feeds mostly on birds, since lemmings are absent from the country. Thus, the population does not exhibit short-term population cycles that are evident in most of the species' geographical range. The population has, however, gone through a major long-term fluctuation in population size. Because of the stability in hunting effort and reliable hunting records since 1958, the total number of adult foxes killed annually can be used as an index of population size (N t ). An index of carrying capacity (K) from population growth data for five separate time blocks during 1958-2007 revealed considerable variation in K and allowed a novel definition of population density in terms of K, or N t /K. Correlation analysis suggested that the reproductive rate was largely determined by the proportion of territorial foxes in the population. Variation in litter size and cub mortality was, on the other hand, related to climatic variation. Thus, Arctic foxes in Iceland engage in typical contest competition but can adapt their territory sizes in response to both temporal and spatial variation in carrying capacity, resulting in surprisingly little variation in litter size.