A circumpolar study unveils a positive non-linear effect of temperature on arctic arthropod availability that may reduce the risk of warming-induced trophic mismatch for breeding shorebirds.

Seasonally abundant arthropods are a crucial food source for many migratory birds that breed in the Arctic. In cold environments, the growth and emergence of arthropods are particularly tied to temperature. Thus, the phenology of arthropods is anticipated to undergo a rapid change in response to a warming climate, potentially leading to a trophic mismatch between migratory insectivorous birds and their prey. Using data from 19 sites spanning a wide temperature gradient from the Subarctic to the High Arctic, we investigated the effects of temperature on the phenology and biomass of arthropods available to shorebirds during their short breeding season at high latitudes. We hypothesized that prolonged exposure to warmer summer temperatures would generate earlier peaks in arthropod biomass, as well as higher peak and seasonal biomass. Across the temperature gradient encompassed by our study sites (>10°C in average summer temperatures), we found a 3-day shift in average peak date for every increment of 80 cumulative thawing degree-days. Interestingly, we found a linear relationship between temperature and arthropod biomass only below temperature thresholds. Higher temperatures were associated with higher peak and seasonal biomass below 106 and 177 cumulative thawing degree-days, respectively, between June 5 and July 15. Beyond these thresholds, no relationship was observed between temperature and arthropod biomass. Our results suggest that prolonged exposure to elevated temperatures can positively influence prey availability for some arctic birds. This positive effect could, in part, stem from changes in arthropod assemblages and may reduce the risk of trophic mismatch.

Vegetation biomass and topography are associated with seasonal habitat selection and fall translocation behavior in Arctic hares.

Habitat selection theory suggests that environmental features selected at coarse scales reveal fundamental factors affecting animal fitness. When these factors vary across seasons, they may lead to large-scale movements, including long-distance seasonal migrations. We analyzed the seasonal habitat selection of 25 satellite-tracked Arctic hares from a population on Ellesmere Island (Nunavut, Canada) that relocated over 100 km in the fall. Since no other lagomorph is known to perform such extensive movements, this population offered an ideal setting to test animal movement and habitat selection theory. On summer grounds hares selected low elevation areas, while on winter grounds they selected high vegetation biomass, high elevation, and steep slopes. During fall relocation, they alternated between stopover and traveling behavioral states (ratio 2:1). Stopover locations were characterized by higher vegetation heterogeneity and lower rugosity than traveling locations, while vegetation biomass and elevation interacted to explain stopover locations in a more complex way. The selected combination of environmental features thus varied across seasons and behavioral states, in a way broadly consistent with predictions based on the changing food and safety needs of hares. Although causality was not demonstrated, our results improve our understanding of long-distance movements and habitat selection in Arctic hares, as well as herbivore ecology in the polar desert. Results also provide strong support to animal movement and habitat selection theory, by showing how some important hypotheses hold when tested in a species phylogenetically distinct from most animal models used in this research field.

Predator-mediated interactions through changes in predator home range size can lead to local prey exclusion.

The strength of indirect biotic interactions is difficult to quantify in the wild and can alter community composition. To investigate whether the presence of a prey species affects the population growth rate of another prey species, we quantified predator-mediated interaction strength using a multi-prey mechanistic model of predation and a population matrix model. Models were parametrized using behavioural, demographic and experimental data from a vertebrate community that includes the arctic fox (Vulpes lagopus), a predator feeding on lemmings and eggs of various species such as sandpipers and geese. We show that the positive effects of the goose colony on sandpiper nesting success (due to reduction of search time for sandpiper nests) were outweighed by the negative effect of an increase in fox density. The fox numerical response was driven by changes in home range size. As a result, the net interaction from the presence of geese was negative and could lead to local exclusion of sandpipers. Our study provides a rare empirically based model that integrates mechanistic multi-species functional responses and behavioural processes underlying the predator numerical response. This is an important step forward in our ability to quantify the consequences of predation for community structure and dynamics.

Predator home range size mediates indirect interactions between prey species in an arctic vertebrate community.

Indirect interactions are widespread among prey species that share a common predator, but the underlying mechanisms driving these interactions are often unclear, and our ability to predict their outcome is limited. Changes in behavioural traits that impact predator space use could be a key proximal mechanism mediating indirect interactions, but there is little empirical evidence of the causes and consequences of such behavioural-numerical response in multispecies systems. Here, we investigate the complex ecological relationships between seven prey species sharing a common predator. We used a path analysis approach on a comprehensive 9-year data set simultaneously tracking predator space use, prey densities and prey mortality rate on key species of a simplified Arctic food web. We show that high availability of a clumped and spatially predictable prey (goose eggs) leads to a twofold reduction in predator (arctic fox) home range size, which increases local predator density and strongly decreases nest survival of an incidental prey (American golden plover). On the contrary, a scattered cyclic prey with potentially lower spatial predictability (lemming) had a weaker effect on fox space use and an overall positive impact on the survival of incidental prey. These contrasting effects underline the importance of studying behavioural responses of predators in multiprey systems and to explicitly integrate behavioural-numerical responses in multispecies predator-prey models.

Warming in the land of the midnight sun: breeding birds may suffer greater heat stress at high- versus low-Arctic sites.

Rising global temperatures are expected to increase reproductive costs for wildlife as greater thermoregulatory demands interfere with reproductive activities. However, predicting the temperatures at which reproductive performance is negatively impacted remains a significant hurdle. Using a thermoregulatory polygon approach, we derived a reproductive threshold temperature for an Arctic songbird-the snow bunting (Plectrophenax nivalis). We defined this threshold as the temperature at which individuals must reduce activity to suboptimal levels (i.e. less than four-time basal metabolic rate) to sustain nestling provisioning and avoid overheating. We then compared this threshold to operative temperatures recorded at high (82° N) and low (64° N) Arctic sites to estimate how heat constraints translate into site-specific impacts on sustained activity level. We predict buntings would become behaviourally constrained at operative temperatures above 11.7°C, whereupon they must reduce provisioning rates to avoid overheating. Low-Arctic sites had larger fluctuations in solar radiation, consistently producing daily periods when operative temperatures exceeded 11.7°C. However, high-latitude birds faced entire, consecutive days when parents would be unable to sustain required provisioning rates. These data indicate that Arctic warming is probably already disrupting the breeding performance of cold-specialist birds and suggests counterintuitive and severe negative impacts of warming at higher latitude breeding locations.

Disentangling the relative influences of global drivers of change in biodiversity: A study of the twentieth-century red fox expansion into the Canadian Arctic.

The poleward range shift of the red fox (Vulpes vulpes) > 1,700 km into the Arctic is one of the most remarkable distribution changes of the early twentieth century. While this expansion threatens a smaller arctic ecological equivalent, the arctic fox (Vulpes lagopus), the case became a textbook example of climate-driven range shifts. We tested this classical climate change hypothesis linked to an important range shift which has attracted little research thus far. We analysed Canadian fur harvest data from the Hudson's Bay Company Archives (14 trading posts; 1926-1950), testing hypotheses based on changes in summer and winter climates. Summer warming might have triggered a bottom-up increase in ecosystem productivity, while winter warming might have lowered thermal stress, both favouring red fox expansion. Additionally, we evaluated the hypothesis that red fox expansion was driven by the appearance of human sedentary sites (n = 110) likely bringing food subsidies into the unproductive tundra. Analysis of red fox expansion chronologies showed that expansion speed was higher during warmer winters. However, the expansions occurred under both cooling and warming trends, being faster during cooler summers in the Baffin Island region. The increasing proportion of red fox in fox fur harvests was best explained by human activity, while generalized linear mixed models also revealed a marginal effect of warmer winters. Generalized additive models confirmed human presence as the most important factor explaining rates of change in the proportion of red fox in fox fur harvests. Using historical ecology, we disentangled the relative influences of climate change and anthropogenic habitat change, two global drivers that transformed arctic biodiversity during the last century and will likely continue to do so during this century. Anthropogenic food subsidies, which constitute stable food sources, facilitated the invasion of the tundra biome by a new mammalian predator and competitor, with long-term consequences that still remain to be understood.

Direct and indirect effects of regional and local climatic factors on trophic interactions in the Arctic tundra.

Climate change can impact ecosystems by reshaping the dynamics of resource exploitation for predators and their prey. Alterations of these pathways could be especially intense in ecosystems characterized by a simple trophic structure and rapid warming trends, such as in the Arctic. However, quantifying the multiple direct and indirect pathways through which climate change is likely to alter trophic interactions and their relative strength remains a challenge. Here, we aim to identify direct and indirect causal mechanisms driven by climate affecting predator-prey interactions of species sharing a tundra food web. We based our study on relationships between one Arctic predator (Arctic fox) and its two main prey - lemmings (preferred prey) and snow geese (alternate prey) - which are exposed to variable local and regional climatic factors across years. We used a combination of models mapping multiple causal links among key variables derived from a long-term dataset (21 years). We obtained several possible scenarios linking regional climate factors (Arctic oscillations) and local temperature and precipitation to the breeding of species. Our results suggest that both regional and local climate factors have direct and indirect impacts on the breeding of foxes and geese. Local climate showed a positive causal link with goose nesting success, while both regional and local climate displayed contrasted effects on the proportion of fox breeding. We found no impact of climate on lemming abundance. We observed positive relationships between lemming, fox and goose reproduction highlighting numerical and functional responses of fox to the variability of lemming abundance. Our study measures causal links and strength of interactions in a food web, quantifying both numerical response of a predator and apparent interactions between its two main prey. These results improve our understanding of the complex effects of climate on predator-prey interactions and our capacity to anticipate food web response to ongoing climate change.

Les changements climatiques peuvent avoir un impact sur les écosystèmes au travers des modifications de la dynamique d'exploitation des ressources par les prédateurs et leurs proies. Dans le cas de l'Arctique, caractérisée par un réseau trophique simple et une sensibilité marquée au réchauffement climatique, l'altération de ces relations trophiques pourrait être particulièrement importante. Cependant, la quantification des nombreux liens directs et indirects à travers lesquels les changements climatiques peuvent affecter les interactions trophiques demeure un défi. Notre objectif est d'identifier les mécanismes causaux directs et indirects, sous-tendus par le climat, affectant les interactions prédateur-proie au sein d'un réseau trophique au cœur de la toundra. Notre étude se base sur les relations entre un prédateur (renard arctique) et ses deux proies principales -le lemming (proie préférée) et la grande oie des neiges (proie alternative)- et qui subissent un accroissement des précipitations et des températures au travers des années. Nous avons utilisé une combinaison de modèles illustrant les liens causaux multiples entre les variables clés issues d'une base de données à long-terme (21 ans). Nous avons obtenu plusieurs scénarios possibles reliant les facteurs climatiques régionaux (Oscillation Arctique) et les températures et précipitations locales à la reproduction de nos 3 espèces. Nos résultats suggèrent que les facteurs climatiques régionaux et locaux présentent des impacts directs et indirects sur la reproduction du renard arctique et de l'oie des neiges. Le climat local présente un lien causal positif avec le succès de nidification de l'oie, alors que le climat local et régional démontrent un effet contrasté sur la proportion de renard en reproduction. Aucune relation entre les facteurs climatiques et l'abondance des lemmings n'a été trouvée. Nous avons observé des liens causaux positifs entre la reproduction du lemming, du renard et de l'oie, mettant en évidence les réponses numériques et fonctionnelles du renard arctique face à la variabilité de l'abondance de lemming. Notre étude est une des premières à mesurer les liens causaux et les forces d'interaction entre les espèces partageant un même réseau trophique, quantifiant ainsi la réponse numérique du prédateur et les interactions apparentes entre ses proies principales. Ces résultats améliorent notre compréhension des effets complexes du climat sur les interactions prédateur-proie et notre capacité à anticiper la réponse des réseaux trophiques aux changements climatiques en cours.

Pulsed food resources affect reproduction but not adult apparent survival in arctic foxes.

As top or mesopredators, carnivores play a key role in food webs. Their survival and reproduction are usually thought to be influenced by prey availability. However, simultaneous monitoring of prey and predators is difficult, making it challenging to evaluate the impacts of prey on carnivores' demography. Using 13 years of field data on arctic foxes Vulpes lagopus in the Canadian High Arctic and a capture-recapture multi-event statistical approach, we investigated the hypothesis that increases in lemming abundance (a cyclic and unpredictable food source) and goose colony proximity (a stable but spatially and temporally limited food source) would be associated with increased apparent survival and reproduction probabilities of adults. Adult apparent survival varied greatly across years (0.13-1.00) but was neither affected by lemming nor goose variations in abundance. However, reproduction probabilities were strongly influenced by both lemming abundance and access to the goose colony. A fox breeding in the best conditions of food availability (year of high lemming density inside the goose colony) had a reproduction probability four times higher than one experiencing the worst conditions (year of low lemming density outside the goose colony). Breeding status of individuals also played a role, with breeders having a 10-20% higher probability of survival and 30% higher probability of reproduction the following year than non-breeders. As the Arctic ecosystem changes due to increased temperatures and species ranges, this study will allow better predictions of predator responses to management or environmental changes and a better understanding of ecosystem functioning.

Seasonal food webs with migrations: multi-season models reveal indirect species interactions in the Canadian Arctic tundra.

Models incorporating seasonality are necessary to fully assess the impact of global warming on Arctic communities. Seasonal migrations are a key component of Arctic food webs that still elude current theories predicting a single community equilibrium. We develop a multi-season model of predator-prey dynamics using a hybrid dynamical systems framework applied to a simplified tundra food web (lemming-fox-goose-owl). Hybrid systems models can accommodate multiple equilibria, which is a basic requirement for modelling food webs whose topology changes with season. We demonstrate that our model can generate multi-annual cycling in lemming dynamics, solely from a combined effect of seasonality and state-dependent behaviour. We compare our multi-season model to a static model of the predator-prey community dynamics and study the interactions between species. Interestingly, including seasonality reveals indirect interactions between migrants and residents not captured by the static model. Further, we find that the direction and magnitude of interactions between two species are not necessarily accurate using only summer time-series. Our study demonstrates the need for the development of multi-season models and provides the tools to analyse them. Integrating seasonality in food web modelling is a vital step to improve predictions about the impacts of climate change on ecosystem functioning. This article is part of the theme issue 'The changing Arctic Ocean: consequences for biological communities, biogeochemical processes and ecosystem functioning'.

Identifying key needs for the integration of social-ecological outcomes in arctic wildlife monitoring.

For effective monitoring in social-ecological systems to meet needs for biodiversity, science, and humans, desired outcomes must be clearly defined and routes from direct to derived outcomes understood. The Arctic is undergoing rapid climatic, ecological, social, and economic changes and requires effective wildlife monitoring to meet diverse stakeholder needs. To identify stakeholder priorities concerning desired outcomes of arctic wildlife monitoring, we conducted in-depth interviews with 29 arctic scientists, policy and decision makers, and representatives of indigenous organizations and nongovernmental organizations. Using qualitative content analysis, we identified and defined desired outcomes and documented links between outcomes. Using network analysis, we investigated the structure of perceived links between desired outcomes. We identified 18 desired outcomes from monitoring and classified them as either driven by monitoring information, monitoring process, or a combination of both. Highly cited outcomes were make decisions, conserve, detect change, disseminate, and secure food. These reflect key foci of arctic monitoring. Infrequently cited outcomes (e.g., govern) were emerging themes. Three modules comprised our outcome network. The modularity highlighted the low strength of perceived links between outcomes that were primarily information driven or more derived (e.g., detect change, make decisions, conserve, or secure food) and outcomes that were primarily process driven or more derived (e.g., cooperate, learn, educate). The outcomes expand monitoring community and disseminate created connections between these modules. Key desired outcomes are widely applicable to social-ecological systems within and outside the Arctic, particularly those with wildlife subsistence economies. Attributes and motivations associated with outcomes can guide development of integrated monitoring goals for biodiversity conservation and human needs. Our results demonstrated the disconnect between information- and process-driven goals and how expansion of the monitoring community and improved integration of monitoring stakeholders will help connect information- and process-derived outcomes for effective ecosystem stewardship.

Identificación de las Necesidades Clave para la Integración de Resultados Socio-Ecológicos en el Monitoreo de Fauna en el Ártico Resumen Para que el monitoreo efectivo en los sistemas socio-ecológicos cumpla con las necesidades de la biodiversidad, la ciencia, y los humanos, se deben definir claramente los resultados deseados y se deben entender las rutas que se toman de los resultados directos hacia los resultados derivados. El Ártico está sufriendo rápidamente cambios climáticos, ecológicos, y económicos, y requiere de un monitoreo efectivo de fauna para cumplir con las necesidades de diversos accionistas. Realizamos entrevistas a profundidad con 29 científicos del Ártico, responsables de decisiones y políticas, y representativos de organizaciones indígenas y organizaciones no gubernamentales para identificar las prioridades de los accionistas con respecto a los resultados deseados del monitoreo de fauna ártica. Mediante un análisis cualitativo de contenido identificamos y definimos los resultados deseados y documentamos las conexiones entre los resultados. Con un análisis de redes investigamos la estructura de las conexiones percibidas y las clasificamos como causadas por el monitoreo de información, el monitoreo del proceso, o una combinación de ambos. Los resultados con un mayor número de menciones fueron tomar decisiones, conservar, detectar cambios, diseminar, y asegurar alimento. Estos reflejan los enfoques más importantes del monitoreo en el Ártico. Los resultados con poca frecuencia en las menciones (p. ej.: regular) correspondían a temas emergentes. Nuestra red de resultados estuvo compuesta por tres módulos. La modularidad resaltó la poca fuerza de las conexiones percibidas entre los resultados que fueron causados principalmente por la información o que estuvieron más derivados (p. ej.: detectar el cambio, tomar decisiones, conservar o asegurar alimento) y los resultados que fueron causados principalmente por el proceso o que estuvieron más derivados (p. ej.: cooperar, aprender, educar). Los resultados expanden la comunidad monitora y diseminan las conexiones creadas entre estos módulos. Los resultados clave deseados se pueden aplicar extensamente a los sistemas socio-ecológicos dentro y fuera del Ártico, particularmente aquellos con economías de sustento basadas en la fauna. Los atributos y motivaciones asociados con los resultados pueden guiar el desarrollo de los objetivos integrados de monitoreo para la conservación de la biodiversidad y las necesidades humanas. Nuestros resultados demostraron la desconexión entre los objetivos conducidos por la información y aquellos conducidos por el proceso y cómo la expansión de la comunidad monitora y una mejor integración de los accionistas monitores ayudarán a conectar los resultados derivados de la información y derivados del proceso para una administración efectiva del ecosistema.

Discrimination factors of carbon and nitrogen stable isotopes from diet to hair in captive large Arctic carnivores of conservation concern.

RATIONALE: Stable isotope analysis is widely used to reconstruct diet, delineate trophic interactions, and determine energy pathways. Such ecological inferences are based on the idea that animals are, isotopically, what they eat but with a predictable difference between the isotopic ratio of a consumer and that of its diet, coined as the discrimination factor. Providing correct estimates of diet-consumer isotopic discrimination in controlled conditions is key for a robust application of the stable isotopes technique in the wild. METHODS: Using a Finnigan Mat Delta Plus isotope-ratio mass spectrometer, we investigated isotopic discrimination of carbon and nitrogen isotope ratios (δ13 C and δ15 N values) in guard hairs of four Arctic predators; the wolf (n = 7), the wolverine (n = 2), the grizzly bear (n = 2), and the polar bear (n = 3). During a 3-month trial, carnivores were fed a mixed diet. The δ13 C and δ15 N values, and the mass (g) of diet items, were monitored weekly for each individual to determine their Total Diet Average ratios. RESULTS: Diet-hair isotopic discrimination (Δx) varied according to species, ranging [1.88 ± 0.69‰: 3.2 ± 0.69‰] for δ13 C values, and [1.58 ± 0.17‰: 3.81 ± 0.22‰] for δ15 N values. Adult wolves Δ13 C average (2.03 ± 0.7‰) was lower than that of young wolves (2.60 ± 0.8‰) and any other species (combined average of 2.59 ± 0.28‰), except for the wolverine (2.12 ± 0.23‰). Wolves Δ15 N averages (juveniles: 3.51 ± 0.34‰, adults: 3.68 ± 0.28‰) were higher than those of any other species (combined average: 2.50 ± 0.58‰). CONCLUSIONS: The discrimination factors for δ13 C and δ15 N values calculated in this study could be used in ecological studies dealing with free-ranging animals, with implications for non-invasive research approaches. As in other controlled discrimination studies, we recommend caution in applying our discrimination factors when the population structure is heterogeneous.

High Arctic lemmings remain reproductively active under predator-induced elevated stress.

Non-consumptive effects of predation have rarely been assessed in wildlife populations even though their impact could be as important as lethal effects. Reproduction of individuals is one of the most important demographic parameters that could be affected by predator-induced stress, which in turn can have important consequences on population dynamics. We studied non-consumptive effects of predation on the reproductive activity (i.e., mating and fertilization) of a cyclic population of brown lemmings exposed to intense summer predation in the Canadian High Arctic. Lemmings were live-trapped, their reproductive activity (i.e., testes visible in males, pregnancy/lactation in females) assessed, and predators were monitored during the summers of 2014 and 2015 within a 9 ha predator-reduction exclosure delimited by a fence and covered by a net, and on an 11 ha control area. Stress levels were quantified non-invasively with fecal corticosterone metabolites (FCM). We found that FCM levels of lemmings captured outside the predator exclosure (n = 50) were 1.6 times higher than inside (n = 51). The proportion of pregnant/lactating adult females did not differ between the two areas, nor did the proportion of adult scrotal males. We found that lemmings showed physiological stress reactions due to high predation risk, but had no sign of reduced mating activity or fertility. Thus, our results do not support the hypothesis of reproductive suppression by predator-induced stress.

Top-down limitation of lemmings revealed by experimental reduction of predators.

It is generally recognized that delayed density-dependence is responsible for cyclic population dynamics. However, it is still uncertain whether a single factor can explain why some rodent populations fluctuate according to a 3-4 yr periodicity. There is increasing evidence that predation may play a role in lemming population cycles, although this effect may vary seasonally. To address this issue, we conducted an experiment where we built a large exclosure (9 ha) to protect brown lemmings (Lemmus trimucronatus) from avian and terrestrial predators. We tested the hypothesis that predation is a limiting factor for lemmings by measuring the demographic consequences of a predator reduction during the growth and peak phases of the cycle. We assessed summer (capture-mark-recapture methods) and winter (winter nest sampling) lemming demography on two grids located on Bylot Island, Nunavut, Canada from 2008 to 2015. The predator exclosure became fully effective in July 2013, allowing us to compare demography between the control and experimental grids before and during the treatment. Lemming abundance, survival and proportion of juveniles were similar between the two grids before the treatment. During the predator-reduction period, summer densities were on average 1.9× higher inside the experimental grid than the control and this effect was greatest for adult females and juveniles (densities 2.4× and 3.4× higher, respectively). Summer survival was 1.6× higher on the experimental grid than the control whereas body mass and proportion of juveniles were also slightly higher. Winter nest densities remained high inside the predator reduction grid following high summer abundance, but declined on the control grid. These results confirm that predation limits lemming population growth during the summer due to its negative impact on survival. However, it is possible that in winter, predation may interact with other factors affecting reproduction and ultimately population cycles.

Cross-scale integration of knowledge for predicting species ranges: a metamodeling framework.

AIM: Current interest in forecasting changes to species ranges have resulted in a multitude of approaches to species distribution models (SDMs). However, most approaches include only a small subset of the available information, and many ignore smaller-scale processes such as growth, fecundity, and dispersal. Furthermore, different approaches often produce divergent predictions with no simple method to reconcile them. Here, we present a flexible framework for integrating models at multiple scales using hierarchical Bayesian methods. LOCATION: Eastern North America (as an example). METHODS: Our framework builds a metamodel that is constrained by the results of multiple sub-models and provides probabilistic estimates of species presence. We applied our approach to a simulated dataset to demonstrate the integration of a correlative SDM with a theoretical model. In a second example, we built an integrated model combining the results of a physiological model with presence-absence data for sugar maple (Acer saccharum), an abundant tree native to eastern North America. RESULTS: For both examples, the integrated models successfully included information from all data sources and substantially improved the characterization of uncertainty. For the second example, the integrated model outperformed the source models with respect to uncertainty when modelling the present range of the species. When projecting into the future, the model provided a consensus view of two models that differed substantially in their predictions. Uncertainty was reduced where the models agreed and was greater where they diverged, providing a more realistic view of the state of knowledge than either source model. MAIN CONCLUSIONS: We conclude by discussing the potential applications of our method and its accessibility to applied ecologists. In ideal cases, our framework can be easily implemented using off-the-shelf software. The framework has wide potential for use in species distribution modelling and can drive better integration of multi-source and multi-scale data into ecological decision-making.

Seasonal demography of a cyclic lemming population in the Canadian Arctic.

1. The causes of cyclical fluctuations in animal populations remain a controversial topic in ecology. Food limitation and predation are two leading hypotheses to explain small mammal population dynamics in northern environments. We documented the seasonal timing of the decline phases and demographic parameters (survival and reproduction) associated with population changes in lemmings, allowing us to evaluate some predictions from these two hypotheses. 2. We studied the demography of brown lemmings (Lemmus trimucronatus), a species showing 3- to 4-year population cycles in the Canadian Arctic, by combining capture-mark-recapture analysis of summer live-trapping with monitoring of winter nests over a 10-year period. We also examined the effects of some weather variables on survival. 3. We found that population declines after a peak occurred between the summer and winter period and not during the winter. During the summer, population growth was driven by change in survival, but not in fecundity or proportion of juveniles, whereas in winter population growth was driven by changes in late summer and winter reproduction. 4. We did not find evidence for direct density dependence on summer demographic parameters, though our analysis was constrained by the paucity of data during the low phase. Body mass, however, was highest in peak years. 5. Weather effects were detected only in early summer when lemming survival was positively related to snow depth at the onset of melt but negatively related to rainfall. 6. Our results show that high mortality causes population declines of lemmings during summer and fall, which suggests that predation is sufficient to cause population crashes, whereas high winter fecundity is the primary factor leading to population irruptions. The positive association between snow depth and early summer survival may be due to the protective cover offered by snow against predators. It is still unclear why reproduction remains low during the low phase.

The effect of snow cover on lemming population cycles in the Canadian high Arctic.

Rising temperatures and changes in the precipitation regime will have a strong impact on the quality of the snow cover in the Arctic. A snow cover of good quality protecting lemmings from cold temperatures and predators is thought to be an important factor for maintaining the cyclic dynamic of their populations in the tundra. We examined if the characteristics of annual fluctuations (amplitude and shape of phases) in brown lemming (Lemmus trimucronatus) density could be determined by snow depth, snow density, sub-nivean temperature and persistence of snow. Using an 18-year time series of brown lemming abundance on Bylot Island in the Canadian Arctic, we tested if snow variables could explain the residual variation between the observed lemming density and the one predicted by models where cyclicity had been accounted for. Our analysis provides support for the hypothesis that snow cover can affect the amplitude and possibly also the periodicity of lemming population cycles in the High Arctic. Summer abundance of brown lemmings was higher following winters with a deep snow cover and a low-density snow pack near the ground but was unaffected by the date of establishment or melting and duration of the snow cover. Two snow variables showed a temporal trend; mean winter snow depth tended to increase and date of establishment of the hiemal threshold occurred earlier over time. These temporal trends, which should be favourable to lemmings, may explain why healthy population cycles have apparently been maintained at our study site contrary to other Arctic sites.

Long-term satellite tracking reveals patterns of long-distance dispersal in juvenile and adult Arctic foxes (Vulpes lagopus).

Long-distance dispersal plays a key role in species distribution and persistence. However, its movement metrics and ecological implications may differ whether it is undertaken by juveniles (natal dispersal) or adults (breeding dispersal). We investigated the influence of life stage on long-distance dispersal in the Arctic fox, an important tundra predator. We fitted 170 individuals with satellite collars during a 13-year study on Bylot Island (Nunavut, Canada), and analysed the tracks of 10 juveniles and 27 adults engaging in long-distance dispersal across the Canadian High Arctic. This behaviour was much more common than expected, especially in juveniles (62.5%, adults: 19.4%). Emigration of juveniles occurred mainly at the end of summer while departure of adults was not synchronized. Juveniles travelled for longer periods and over longer cumulative distances than adults, but spent similar proportions of their time travelling on sea ice versus land. Successful immigration occurred mostly in late spring and was similar for juveniles and adults (30% versus 37%). Our results reveal how life stage influences key aspects of long-distance dispersal in a highly mobile canid. This new knowledge is critical to understand the circumpolar genetic structure of the species, and how Arctic foxes can spread zoonoses across vast geographical areas.

Born in the cold: contrasted thermal exchanges and maintenance costs in juvenile and adult snow buntings on their breeding and wintering grounds.

Several species of passerines leave their nest with unfinished feather growth, resulting in lower feather insulation and increased thermoregulatory demands compared to adults. However, feather insulation is essential for avian species breeding at northern latitudes, where cold conditions or even snowstorms can occur during the breeding season. In altricial arctic species, increased heat loss caused by poor feather insulation during growth could be counter-adaptative as it creates additional energy demands for thermoregulation. Using flow-through respirometry, we compared resting metabolic rate at thermoneutrality (RMRt), summit metabolic rate (Msum) and heat loss (conductance) in adult and juvenile snow buntings on their summer and winter grounds. In summer, when buntings are in the Arctic, juveniles had a 12% higher RMRt, likely due to unfinished growth, and lost 14% more heat to the environment than adults. This pattern may result from juveniles fledging early to avoid predation at the cost of lower feather insulation. Surprisingly, an opposite pattern was observed at lower latitudes on their wintering grounds. Although they showed no difference in RMRt and Msum, adults were losing 12% more heat than juveniles. We suggest that this difference is due to poorer insulative property of plumage in adults stemming from energetic and time constraints encountered during their post-breeding molt. High plumage insulation in first-winter juvenile buntings could be adaptive to reduce thermoregulatory demands and maximize survival in the first winter of life, while adults could use behavioral strategies to compensate for their greater rate of heat loss.

Benefiting from a migratory prey: spatio-temporal patterns in allochthonous subsidization of an Arctic predator.

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

Long-distance, synchronized and directional fall movements suggest migration in Arctic hares on Ellesmere Island (Canada).

Animal migration contributes largely to the seasonal dynamics of High Arctic ecosystems, linking distant habitats and impacting ecosystem structure and function. In polar deserts, Arctic hares are abundant herbivores and important components of food webs. Their annual migrations have long been suspected, but never confirmed. We tracked 25 individuals with Argos satellite telemetry to investigate the existence of migration in a population living at Alert (Ellesmere Island, Nunavut, Canada). During fall, 21 hares undertook directional, long-distance movements in a southwestern direction towards Lake Hazen. Daily movement rates averaged 1.3 ± 0.5 km, 4.3 ± 1.6 km, and 1.7 ± 0.9 km before, during, and after relocation, respectively. Straight-line and minimum cumulative distances traveled averaged 98 ± 18 km (range: 72-148 km) and 198 ± 62 km (range: 113-388 km), respectively. This is the first report of large-scale seasonal movements in Arctic hares and, surprisingly, in any lagomorph species. These movements may be part of an annual migratory pattern. Our results redefine our understanding of the spatial ecology of Arctic hares, demonstrate unsuspected mobility capacities in lagomorphs, and open new perspectives regarding the ecological dynamics of the northern polar deserts.