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During the cold arctic winter, small mammals like lemmings seek refuge inside the snowpack to keep warm and they dig tunnels in the basal snow layer, usually formed of a soft depth hoar, to find vegetation on which they feed. The snowpack, however, is a heterogenous medium and lemmings should use habitats where snow properties favor their survival and winter reproduction. We determined the impact of snow physical properties on lemming habitat use and reproduction in winter by sampling their winter nests for 13 years and snow properties for 6 years across 4 different habitats (mesic, riparian, shrubland, and wetland) on Bylot Island in the Canadian High Arctic. We found that lemmings use riparian habitat most intensively because snow accumulates more rapidly, the snowpack is the deepest and temperature of the basal snow layer is the highest in this habitat. However, in the deepest snowpacks, the basal depth hoar layer was denser and less developed than in habitats with shallower snowpacks, and those conditions were negatively related to lemming reproduction in winter. Shrubland appeared a habitat of moderate quality for lemmings as it favored a soft basal snow layer and a deep snowpack compared with mesic and wetland, but snow conditions in this habitat critically depend on weather conditions at the beginning of the winter. With climate change, a hardening of the basal layer of the snowpack and a delay in snow accumulation are expected, which could negatively affect the winter habitat of lemmings and be detrimental to their populations., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Highly mobile predators can show strong numerical responses to pulsed resources, sometimes resulting in irruptions where large numbers of young invade landscapes at a continental scale. High production of young in irruption years may have a strong influence on the population dynamics unless immature survival is reduced compared to non-irruption years. This could occur if subordinate individuals (mainly immatures) are forced into suboptimal habitats due to density-dependent effects in irruption years. To test whether irruptive individuals had lower survival than non-irruptive ones, we combined necropsy results (N = 365) with telemetry (N = 185) from more than 20 years to record timing and causes of mortality in snowy owls (Bubo scandiacus), which irrupt into eastern North America during winter following high breeding output caused by lemming peaks in the Arctic. Mortality was more than four times higher in irruption years than non-irruption years, but only for immatures, and occurred disproportionately in early winter for immatures, but not adults. Mortality was also higher in eastern North America, where owl abundance fluctuates considerably between years, compared to core winter regions of the Arctic and Prairies where populations are more stable. Most mortality was not due to starvation, but rather associated with human activity, especially vehicle collisions. We conclude that immature snowy owls that irrupt into eastern North America are limited by density-dependent factors, such as increased competition forcing individuals to occupy risky human-altered habitats. For highly mobile, irruptive animals, resource pulses may have a limited impact on population dynamics due to low subsequent survival of breeding output during the nonbreeding season., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

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.

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.

Studies of population dynamics of long-lived species have generally focused on adult survival because population growth should be most sensitive to this parameter. However, actual variations in population size can often be driven by other demographic parameters, such as juvenile survival, when they show high temporal variability. We used capture-recapture data from a long-term study of a hunted, migratory species, the greater snow goose (Chen caerulescens atlantica), to assess temporal variability in first-year survival and the relative importance of natural and hunting mortality. We also conducted a parasite-removal experiment to determine the effect of internal parasites and body condition on temporal variation in juvenile survival. We found that juvenile survival showed a higher temporal variability than adult survival and that natural mortality was more important than hunting mortality, unlike in adults. Parasite removal increased first-year survival and reduced its annual variability in females only. Body condition at fledging was also positively correlated with first-year survival in treated females. With reduced parasite load, females, which are thought to invest more in their immune system than males according to Bateman's principle, could probably reallocate more energy to growth than males, leading to a higher survival. Treated birds also had a higher survival than control ones during their second year, suggesting a developmental effect that manifested later in life. Our study shows that natural factors such as internal parasites may be a major source of variation in juvenile survival of a long-lived, migratory bird, which has implications for its population dynamics.

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.

The insulative value of early and deep winter snow is thought to enhance winter reproduction and survival by arctic lemmings (Lemmus and Dicrostonyx spp). This leads to the general hypothesis that landscapes with persistently low lemming population densities, or low amplitude population fluctuations, have a low proportion of the land base with deep snow. We experimentally tested a component of this hypothesis, that snow depth influences habitat choice, at three Canadian Arctic sites: Bylot Island, Nunavut; Herschel Island, Yukon; Komakuk Beach, Yukon. We used snow fencing to enhance snow depth on 9-ha tundra habitats, and measured the intensity of winter use of these and control areas by counting rodent winter nests in spring. At all three sites, the density of winter nests increased in treated areas compared to control areas after the treatment, and remained higher on treated areas during the treatment. The treatment was relaxed at one site, and winter nest density returned to pre-treatment levels. The rodents' proportional use of treated areas compared to adjacent control areas increased and remained higher during the treatment. At two of three sites, lemmings and voles showed significant attraction to the areas of deepest snow accumulation closest to the fences. The strength of the treatment effect appeared to depend on how quickly the ground level temperature regime became stable in autumn, coincident with snow depths near the hiemal threshold. Our results provide strong support for the hypothesis that snow depth is a primary determinant of winter habitat choice by tundra lemmings and voles.

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.

Despite numerous studies on breeding dispersal, it is still unclear how habitat heterogeneity and previous nesting success interact to determine nest-site fidelity at various spatial scales. In this context, we investigated factors affecting breeding dispersal in greater snow geese (Anser caerulescens atlanticus), an Arctic breeding species nesting in two contrasting habitats (wetlands and mesic tundra) with variable pattern of snowmelt at the time of settlement in spring. From 1994 to 2005, we monitored the nesting success and breeding dispersal of individually marked females. We found that snow geese showed a moderate amount of nest-site fidelity and considerable individual variability in dispersal distance over consecutive nesting attempts. This variability can be partly accounted for by the annual timing of snowmelt. Despite this environmental constraint, habitat differences at the colony level consistently affected nesting success and settlement patterns. Females nesting in wetlands had higher nesting success than those nesting in mesic tundra. Moreover, geese responded adaptively to spatial heterogeneity by showing fidelity to their nesting habitat, independently of snowmelt pattern. From year to year, geese were more likely to move from mesic to high-quality wetland habitat, regardless of previous nesting success and without cost on their subsequent nesting performance. The unpredictability of snowmelt and the low cost of changing site apparently favour breeding-site dispersal although habitat quality promotes fidelity at the scale of habitat patches.

Even though growth rate is an important fitness component, it is still controversial to what extent parent birds adjust the timing of offspring hatch to natural variations in food supply to maximize offspring growth. We studied the role of food availability in explaining inter- and intra-seasonal variation of growth rate in goslings of greater snow geese over 5 years. The peak of hatching generally coincided with the peak of food availability. However, early-hatched goslings usually grew faster than birds hatched at the peak, which in␣turn grew faster than late-hatched goslings, although this phenomenon was not observed in all years. There was considerable variation in growth rate among the five years, the smallest goslings produced in the best year (1991) being larger than the largest goslings of the poorest year (1994). We developed three indices of food availability, based on the cumulative availability of plant biomass and nitrogen content during the growth period, and showed that the cumulative exposure to nitrogen biomass explained up to 43% of variation (intra- and inter-annual) in body size just before fledging. In years with good feeding conditions, early-hatched goslings had access to more nitrogen during their growing period than those hatching on or after the peak and they grew faster. In years of lower food availability, early-hatched goslings had no detectable advantage over peak- or late-hatched birds for access to protein-rich food and no seasonal decline in growth rate was observed. These results confirm the critical role of food supply in the seasonal variation of growth rate in Arctic-nesting geese.

Temporary feeding on willow buds and leaves by nesting greater snow geese provided us with an opportunity to test the relative importance of nutrients and deterrents in affecting the palatability for geese of a food plant with a high phenol content. Protein, total phenol and fiber (neutral and acid detergent fiber, and lignin) were analyzed in closed and open buds and in rolled and open leaves. Geese feed on willows at the open-buds and rolled-leaf stages but not at the closed-bud and open-leaf stages. Protein content was higher in open buds and rolled leaves (25-27%) than in closed buds and open leaves (19-21%). Phenol content increased during leaf emergence but was already high (14%) in rolled leaves. All plant fibers were very high in closed buds but declined rapidly during leaf emergence. The increase in phenol: protein ratio appeared to be more important than phenol concentration alone in explaining the cessation of feeding by geese on willow leaves whereas the high fiber content of closed buds may explain why they were not eaten. Our results illustrate the value of a multifactorial approach in the study of the food selection process in herbivores.

Arctic-nesting geese are classified as "capital" breeders (i.e., birds that rely largely on endogenous reserves to meet the high nutrient requirement of clutch formation) as opposed to "income" breeders (those that rely directly on ingested food). However, some evidence has suggested that energy reserves of greater snow geese (Chen caerulescens atlantica) are not sufficient to meet these costs. We tested whether greater snow geese were capital breeders and examined the consequences on their reproductive strategy. We collected 48 females and 47 males from arrival at the breeding colony on Bylot Island (NWT) to the postlaying stage, and determined fat and protein content in somatic and reproductive tissues. Rapid follicular growth was initiated only after arrival on the breeding ground. Somatic fat of females tended to increase during the prelaying and laying stages at the same time that most of the fat was deposited in developing follicles. Decrease in somatic fat in postlaying females was mostly related to its use for meeting metabolic requirements during early incubation. Hence, almost all fat invested in the clutch came from food intake, not endogenous reserves. Somatic protein was maintained during prelaying but decreased during laying, suggesting that some protein reserves were deposited in the eggs. There was no relationship between somatic fat and the number of developing follicles (incubating females excluded) but a relationship was found with somatic protein. Clutch size was not related to body size. In males, somatic fat, which was similar to females at arrival, was almost completely depleted by the postlaying stage. Male somatic protein remained stable. Male somatic reserves were not related to the somatic reserves, clutch size or nutrient investment in reproduction of their mate. We conclude that, in female greater snow geese, little fat reserves (if any) were used for egg formation though some protein reserves were used, whereas males relied heavily on their fat reserves during the same period. This is probably a consequence of the long and costly migration between the last staging area and the breeding grounds.

We tested Buchsbaum's hypothesis that food palatability in geese is determined by a hierarchy of feeding cues among which deterrent secondary metabolites (mostly phenols) have a primary role (Buchsbaum et al. 1984). In preference tests, greater snow goose feeding was slightly depressed when grass was sprayed with ferulic acid but not when grass was sprayed with p-coumaric and tannic acids. Extracts of Timothy grass, red clover or alfalfa sprayed on grass also failed to depress goose feeding. In a multifactor experiment, phenol and protein content and height of grass were manipulated simultaneously. When ferulic acid was sprayed, protein and phenol content interacted in determining goose feeding preferences; protein content had no effect in the absence of phenol but did have an effect when phenol was added. When tannic acid was used in a similar experiment, results were inconclusive because of a significant and complex interaction between protein content and height of grass. Our results generally failed to support Buchsbaum's hypothesis that phenol content of plants has a primary role in determining food preference in geese. Protein content of plants seemed to be a more important factor.