Population limitation of the northern red-backed vole in the boreal forests of northern Canada

1. Across the vast boreal forests of North America, no population cycles in Clethrionomys species occur. In Eurasia, by contrast, some Clethrionomys populations of the same species undergo regular 3-5-year cycles. We examined the effects of nutrients, food, competitors, predators and climate on population limitation in the northern red-backed vole (Clethrionomys rutilus Pallas) in the south-western Yukon to determine why this difference occurs. 2. From 1986 to 1996 we added food, reduced large mammal predators and excluded snowshoe hares (Lepus americanus Erxleben) from large plots and found that none of these manipulations affected red-backed vole abundance. Adding nutrients as nitrogen, phosphorus and potassium (NPK) fertilizer had a slight negative effect, probably acting through a reduction in dwarf shrub productivity caused by competition from grasses. 3. We monitored weasel populations directly through trapping and indirectly through snow tracking. Predation by these vole specialists was irrelevant as a limiting factor most of the time because voles in this area do not reach the densities needed to sustain weasel populations. Other boreal forest mammal and bird predators did not focus on red-backed voles. However, when red-backed vole populations increased in the forest and Microtus voles also increased in the meadows, weasel populations increased and may have temporarily depressed red-backed voles in winter. 4. We monitored one major potential food, white spruce seeds, but seed fall was not related to population changes in red-backed voles, even after mast years. 5. We assessed the impact of weather variables, and the average depth of the snow pack during winter (October-March) was correlated directly with vole demography, having both direct effects in that year and delayed effects in the following year. 6. Our long-term trapping data (1973-96) indicate that Clethrionomys populations fluctuated, with peaks following hare peaks by 2-3 years. 7. We propose that the key variable limiting these vole populations is overwinter survival, and this is a function of overwinter food from berries produced during the previous summer by dwarf shrubs. These shrubs may be stimulated by abundant moisture from winter snows or by periodic fertilization from large quantities of pellets produced at snowshoe hare peaks.     

From process to pattern: how fluctuating predation risk impacts the stress axis of snowshoe hares during the 10-year cycle

Predation is a central organizing process affecting populations and communities. Traditionally, ecologists have focused on the direct effects of predation—the killing of prey. However, predators also have significant sublethal effects on prey populations. We investigated how fluctuating predation risk affected the stress physiology of a cyclic population of snowshoe hares (Lepus americanus) in the Yukon, finding that they are extremely sensitive to the fluctuating risk of predation. In years of high predator numbers, hares had greater plasma cortisol levels at capture, greater fecal cortisol metabolite levels, a greater plasma cortisol response to a hormone challenge, a greater ability to mobilize energy and poorer body condition. These indices of stress had the same pattern within years, during the winter and over the breeding season when the hare:lynx ratio was lowest and the food availability the worst. Previously we have shown that predator-induced maternal stress lowers reproduction and compromises offspring’s stress axis. We propose that predator-induced changes in hare stress physiology affect their demography through negative impacts on reproduction and that the low phase of cyclic populations may be the result of predator-induced maternal stress reducing the fitness of progeny. The hare population cycle has far reaching ramifications on predators, alternate prey, and vegetation. Thus, predation is the predominant organizing process for much of the North American boreal forest community, with its indirect signature—stress in hares—producing a pattern of hormonal changes that provides a sensitive reflection of fluctuating predator pressure that may have long-term demographic consequences.     

Overwinter mass loss of snowshoe hares in the Yukon: starvation, stress, adaptation or artefact?

1. Overwinter mass loss can reduce energetic requirements in mammals (Dehnel's phenomenon). Alternatively, mass loss can result from food limitation or high predation risk. 2. We use data from fertilizer, food-supplementation and predator-exclusion experiments in the Yukon during a population cycle from 1986 to 1996 to test the causes of overwinter mass loss by snowshoe hares (Lepus americanus). In all years, some hares on control sites gained mass overwinter. During the increase phase the majority gained mass, but in all other phases the majority lost mass. 3. Snowshoe hares weighing <1000 g in autumn always gained mass overwinter, as did the majority that weighed 1000-1400 g. Hares weighing >1800 g in autumn usually lost mass. 4. Snowshoe hares on the predator-exclosure + food site gained mass overwinter in all years. Hares on the food-supplementation sites lost mass during the decline but gained mass in all other phases. Fertilization had little effect on mass dynamics. 5. Snowshoe hares were more likely to lose mass during winters with low survival rates. Snowshoe hares on the predator-exclosure treatments were more likely to gain mass than were hares on control sites. 6. Overwinter mass loss was correlated with maximum snow depth. At equivalent snow depths, hares on food-supplemented areas lost 98 g (+/- 14.6 SE) less on average than hares on the controls and predator-exclosure treatment. 7. Bone-marrow fat was related to body mass and cause of death. Small hares had the lowest marrow fat. Hares killed by humans had higher marrow fat than those killed by predators; hares that simply died had the lowest marrow fat. Hares on food-supplemented sites had the highest kidney and marrow fat. 8. Overwinter-mass loss for snowshoe hares is explained interactively by winter conditions, food supply, predation risk and autumn mass. Some snowshoe hares lost mass overwinter in all years and on all treatments, suggesting that reducing body mass may facilitate survival, especially in cases where foraging costs are high energetically or increase predation risk.     

Snowshoe hare demography during a cyclic population low

1. Snowshoe hare ( Lepus americanus Erxleben) populations were studied in south-west Yukon during the low phase of the 10-year population cycle. Food availability and predator abundance were manipulated in a factorial design to determine the importance of each factor in hare dynamics during this phase.
2. Food was abundant during the low phase, and snowshoe hares were not food limited.
3. Survival of hares was higher than at any other phase of the cycle, and predators were scarce, but >75% of hare deaths resulted from predation.
4. Food addition resulted in higher hare densities and better body condition than on control sites. There were no observable effects of food addition on population rate of increase, recruitment, survival or age structure.
5. Mammalian predator reduction resulted in higher hare densities, higher survival, better body condition and an older age structure. Relative to control populations, recruitment was lower and population rates of increase similar.
6. The joint manipulation of food addition + predator reduction had greater positive effects on hare density and body condition than either single factor manipulation. Survival was better than on control sites, and the age structure was older than on control sites. Population rates of increase were similar, but recruitment was higher on the control areas.
7. We conclude that snowshoe hare dynamics at the low of the cycle are dominated by the interaction of food and predation. Risk of predation also had indirect effects on snowshoe hare age structure and body condition.     

Effect of late winter food addition on numbers and movements of snowshoe hares

Summary Peak (1980) and early decline (1981) populations of snowshoe hares were supplied with extra food in late winter to test the hypothesis that snowshoe hare populations are limited by food supply. Food supplemented populations increased through immigration in both years but the response was more pronounced in the early decline population. Animals supplied with extra food lost less weight, had higher survival in some cases, and males began to breed earlier. Immigrants to the food addition area were of two types: those that established home ranges on or near the area and those that spent only a brief time there before returning to their initial range. The possibility that these latter individuals were prevented from remaining on the food grid by residents is discussed. Results indicate that food supply is one factor that can limit peak and declining populations of snowshoe hares but the relation of spacing behaviour to food supply and numbers must also be considered.     

Unstable dynamics and population limitation in mountain hares

The regular large-scale population fluctuations that characterize many species of northern vertebrates have fascinated ecologists since the time of Charles Elton. There is still, however, no clear consensus on what drives these fluctuations. Throughout their circumpolar distribution, mountain hares Lepus timidus show regular and at times dramatic changes in density. There are distinct differences in the nature, amplitude and periodicity of these fluctuations between regions and the reasons for these population fluctuations and the geographic differences remain largely unknown. In this review we synthesize knowledge on the factors that limit or regulate mountain hare populations across their range in an attempt to identify the drivers of unstable dynamics. Current knowledge of mountain hare population dynamics indicates that trophic interactions--either predator-prey or host-parasite--appear to be the major factor limiting populations and these interactions may contribute to the observed unstable dynamics. There is correlative and experimental evidence that some mountain hare populations in Fennoscandia are limited by predation and that predation may link hare and grouse cycles to microtine cycles. Predation is unlikely to be important in mountain hare populations in Scotland as most hares occur on sporting estates where predators are controlled, but this hypothesis remains to be experimentally tested. There is, however, emerging experimental evidence that some Scottish mountain hare populations are limited by parasites and that host-parasite interactions contribute to unstable dynamics. By contrast, there is little evidence from Fennoscandia that parasitism is of any importance to mountain hare population dynamics, although disease may cause periodic declines. Although severe weather and food limitation may interact to cause periodic high winter mortality there is little evidence that food availability limits mountain hare populations. There is a paucity of information concerning the factors limiting or regulating mountain hare populations in the Alps of Central Europe or in the tundra and taiga belts of Russia. Future research on mountain hare population dynamics should focus on the interactions between predation, parasitism and nutrition with stochastic factors such as climate and anthropogenic management including harvesting.     

ASSESSMENT OF POTENTIAL BIAS WITH SNOWSHOE HARE FECAL PELLET-PLOT COUNTS

Abstract: The fecal pellet-plot method has been used extensively for snowshoe hare (Lepus americanus) population studies across the species' range, but potential biases associated with the technique have not been addressed adequately. We studied hare pellet-plots in northern Idaho to quantify pellet decomposition rates across environmental gradients, and conducted feeding trials on captive hares to assess the role of diet on pellet production rates. We found that across our study area pellet numbers tended to be higher on plots with high vegetative cover, which likely was a reflection of hare habitat choice rather than lesser pellet decomposition in such habitat. A pellet decomposition experiment indicated that pellet persistence was negatively related to moisture level, and that pellets produced by hares during summer decomposed more quickly than those from winter. We found that only 19% of fecal pellets collected from plots located across northern Idaho were produced by hares during winter. There was a correlation between pellet numbers from plots that were pre-cleared 1 year earlier and estimated numbers of hares on 6 study areas. A similar correlation was lacking for pellet counts from uncleared plots, implying that hare population estimation via pellet-plot counts should involve plot pre-clearing. In captive studies, juvenile hares produced slightly fewer pellets per day per gram of food ingested than adults, but pellet production was similar across diets comprised of 10 different browse species. We conclude that for our study area the fecal pellet-plot method may be subject to notable pellet decomposition bias, and therefore recommend that use of the method elsewhere across the species' range be preceded by assessment of both the pellet-hare density relationship and pellet decomposition rates across habitats.     

An examination of a compensatory relationship between food limitation and predation in semi-domestic reindeer

A central issue in ecology is to what extent food limitation and predation affect animal populations. We studied how survival and reproductive success was related to the female's size in a population of semi-domesticated reindeer during 2 years where there was a large difference in snowfall during winter. The females were kept within a predator-free enclosure for about 5 weeks during the calving period and thereafter released to their natural summer pastures. Small females were more likely to fail to reproduce and they produced smaller calves than large females. Additionally, small females were more likely to loose their calves due to starvation within the predator-free enclosure and to predators outside the enclosure. Food limitation during the harsh winter appeared to be the major cause of deaths. However, food limitation interacted with predation and led to high calf losses when the females experienced low food availability during the harsh winter. In contrast, predators killed no calves after the mild winter. Apparently, the interaction between predation and food limitation is due to small females favouring their own growth and survival over calf production in summers following harsh winters with food shortage. Our results indicate that a compensatory relationship exists between mortality due food limitation and predation. Thus, the impact of calf predation on reindeer demography and population dynamics may be limited.     

Mountain hare populations on islands: effects of predation by red fox

Summary On islands off the west coast of Sweden the density of mountain hares (Lepus timidus L.) is very high. One of the main predators on hares, the red fox (Vulpes vulpes L.), is only present during short periods. Data on hare density and predation by red fox and eagle owl (Bubo bubo (L.)) has been analyzed from five islands over several years. Winter mortality in years with low predation pressure was independent of hare density. But when red fox or eagle owl were present on islands (i.e., high predation pressure) winter mortality became density dependent. Thus, at low density, winter mortality did not increase through red fox predation. But at densities up to two hares/ha, predation pressure was increasing and could be limiting for these populations. At still higher hare density predation pressure became less intensive. The functional response for foxes preying on hares showed a type II or a sigmoid type III response pattern. In normal summers, the population increase due to reproduction was at least two-fold. When a fox was present there was instead a sharp decrease in hare numbers. Fox predation had a stronger effect in summer than in winter. By switching between islands and mainland areas from winter to summer, a fox can stabilize fluctuations in hare numbers on the islands. This is dependent on how often the ice permits a fox to reach an island and the lack of numerical response by predators.     

Evidence of autumn reproduction in female European hares (Lepus europaeus) from southern Europe

Information on reproductive biology of the European hare (Lepus europaeus) in different environmental and landscape conditions comprises part of fundamental knowledge regarding species’ adaptive responses as well as many aspects of its biology. Most of the studies conducted on European hare reproduction are confined to midlatitude and northern populations, whereas no data exist on the indigenous southern populations. Here, we present information on reproductive characteristics of European hares inhabiting Mediterranean ecosystems on the island of Crete, Greece for two successive hunting seasons. Although the annual reproductive cycle of the species is well known, with an autumn sexual inactivity, the duration of this period is subjected to fluctuations in different years and for different areas. According to our data, hare populations of Crete present an autumn–early winter reproductive activity with high proportions of pregnant females observed in all the months of the study. Furthermore, the estimated mean litter size (1.54 SE ± 0.07) while signed to the lowest values ever observed for European hares is similar to values obtained in continuous breeding species of the same genus, Lepus granatensis, Lepus corsicanus, Lepus (capensis) mediterraneus, and Lepus capensis also inhabiting warm climates. In conclusion, our results suggest that Cretan European hare populations exhibit a reproductively active period during autumn–early winter where proportions of pregnant females and litter size give a strong indication of a continuous reproduction throughout the year.     

Diet selection by hares (Lepus europaeus) in arable land and its implications for habitat management

Populations of European hares (Lepus europaeus) have experienced a dramatic decline throughout Europe in recent decades. European hares are assumed to prefer weeds over arable crops, and weed abundance was reduced by the intensification of agriculture. Therefore, modern agriculture has been blamed as a major factor affecting European hare populations. However, it is questionable whether European hares select weeds at all, as previous studies had major methodological limitations. By comparing availability and use of plants with Chesson’s Electivity Index, we investigated whether the European hare actually feeds selectively on different plants in arable land. Food availability and use were dominated by cultivated crops (e.g. winter wheat, spring barley and sugar beet). Diet selection analysis revealed that in autumn and winter, European hares predominantly preferred cultivated crops (winter wheat) and food items provided by hunters (tubers of sugar beet and carrot). In spring and summer, apart from soy, only weeds (e.g. clover and corn poppy) were positively selected, especially after cereal crops were harvested. We suggest that the decline in European hare populations throughout Europe was facilitated by the decrease in weed abundance. Wildlife-friendly set-asides in arable land have the potential to reconcile the European Union’s Common Agricultural Policy with wildlife conservation.     

Of lemmings and snowshoe hares: the ecology of northern Canada

Two population oscillations dominate terrestrial community dynamics in northern Canada. In the boreal forest, the snowshoe hare (Lepus americanus) fluctuates in cycles with an 8-10 year periodicity and in tundra regions lemmings typically fluctuate in cycles with a 3-4 year periodicity. I review 60 years of research that has uncovered many of the causes of these population cycles, outline areas of controversy that remain and suggest key questions to address. Lemmings are keystone herbivores in tundra ecosystems because they are a key food resource for many avian and mammalian predators and are a major consumer of plant production. There remains much controversy over the role of predation, food shortage and social interactions in causing lemming cycles. Predation is well documented as a significant mortality factor limiting numbers. Food shortage is less likely to be a major limiting factor on population growth in lemmings. Social interactions might play a critical role in reducing the rate of population growth as lemming density rises. Snowshoe hares across the boreal forest are a key food for many predators and their cycles have been the subject of large-scale field experiments that have pinpointed predation as the key limiting factor causing these fluctuations. Predators kill hares directly and indirectly stress them by unsuccessful pursuits. Stress reduces the reproductive rate of female hares and is transmitted to their offspring who also suffer reduced reproductive rates. The maternal effects produced by predation risk induce a time lag in the response of hare reproductive rate to density, aiding the cyclic dynamics.     

A natural feeding experiment on a declining snowshoe hare population

Summary A snowshoe hare (Lepus americanus) population on a 9-ha area was supplied with extra natural food by chopping down large white spruce (Picea glauca) and aspen (Populus tremuloides) trees throughout 3 winters from 1981 to 1984. Hares fed vigorously on the downed trees, but the phase of decline of the ten-year cycle occurred equally on control and experimental grids from 1981–1983, and we could detect no improvement in survival or reproduction on the food area. Growth rates were improved on the food grid during the first winter of the decline (1981–82). We concluded that food shortage is not necessary for the cyclic decline of snowshoe hares in the southern Yukon.     

The recent expansion of the brown hare (<Emphasis Type="Italic">Lepus europaeus</Emphasis>) in Sweden with possible implications to the mountain hare (<Emphasis Type="Italic">L. timidus</Emphasis>)

The brown hare (Lepus europaeus) expanded its Swedish distribution since the 1980s northwards and locally to new areas within its former range. Of 115 brown hare populations within the former range reported in a hunter enquiry, those established after 1980 were situated higher above the sea level than older ones and higher than neighbouring (<50 km) older populations. Reports on increased use of forest habitats by brown hares were equally frequent among recent and older populations, suggesting a process promoted solely by less harsh winters. Supposed hare hybrids were more often reported from hunting grounds with recent brown hare establishment, i.e. where the species expands in time and in space. In a 27-year dataset on brown hare observations, the recent increased use of forest habitats was supported in that maximum distances to agricultural land for brown hare sightings were higher in mild winters, whereas the proportions of the annual observations made during winter were lower. In 40-year bag records from two Swedish counties, the dynamics of the mountain hare (Lepus timidus) responded positively to snow parameters, whereas brown hares responded negatively. We suggest that the state of mountain hare populations primarily depends on winter conditions and predation pressure, whereas possible effects of hybridization are unclear. If winter conditions remain as in the last 15 years, mountain hare numbers are not likely to increase in southern Sweden, whereas the brown hare may expand even further. In either case, hybrids will occur in sympatric areas in frequencies probably related to the density of the respective true species.     

Winter diet, habitat selection and fluctuation of a mountain hare Lepus timidus population in Finnish Forest Lapland

Composition of the winter diet, habitat selection and population fluctuations in the mountain hare Lepus timidus were studied in the Värriötunturi fell area, Eastern Finnish Forest Lapland, during the winters 1968/69–1984/85. The three population lows recorded during this 17-year period followed each other at intervals of 4 and 8 years. During the lows the hares occurred only in the most favoured (forest-covered) habitats and in two of them they behaved according to the concept of the refuge theory. The mountain birch Betula pubescens ssp. tortuosa appeared to be the most important, but not the most favoured winter food item. When the population crashed, the proportion of birch in the diet decreased, and was replaced especially by juniper which is one of the secondary food items (and is for this reason often discarded although cut). It is suggested that the quality and/or quantity of the winter food (i.e. winter pastures) are one of the driving forces in the population fluctuation of the mountain hare in this area.     

Selection of resting sites by the European hare as related to habitat characteristics during agricultural changes

The decline of the European hare Lepus europaeus populations has been shown to be correlated with agricultural intensification, which has caused loss of habitat heterogeneity. We aimed to investigate the diurnal spring habitat selection using Jacobs’ second selection index in an intensively cultivated farmland to reveal how the habitat choice by resting hares was affected by changes in the habitat availability. Clearance counts of hares driven out of known areas were made during an 11-year field investigation within two sectors of a private hunting ground located in the Paris basin (France). The loss of habitat heterogeneity included the disappearance of pastures dedicated to a last remaining sheep farm in the area, the removal of non-cropped areas and field boundaries and the increase of mean field size. Breeding stocks of hare declined during the study. Harrowed fields were always avoided. Pastures, alfalfa fields, thickets and fallow land were selected at the start of our study, whereas the remaining part was avoided at the end. The preference for ploughed fields decreased with the growth of vegetation in the winter-wheat fields. To benefit hares, land management should provide year-round vegetative cover and food from non-cropped areas in intensive arable farms.     

Parasitism in a declining population of snowshoe hares

Prevalence and intensity of six endoparasites were determined in 346 snowshoe hares (Lepus americanus) obtained at Rochester, Alberta, during December-April 1981-1982, the second winter of a cyclic population decline. The data were analyzed for (1) differences among host sex and age classes, and among months and sample sources, and (2) evidence that parasitism was of demographic significance to the hare population. Prevalence and intensity of Obeliscoides cuniculi were consistently highest among adult hares, but rose most sharply from February to March among juveniles. In contrast, prevalence and intensity of Nematodirus triangularis were highest among juveniles; prevalence reached 90-100% by January, whereas intensity continued to rise through April. Prevalence and intensity of both Trichuris leporis and Protostrongylus boughtoni were highest also among juvenile hares; neither parameter exhibited a definite trend over time. Prevalences of Taenia pisiformis (cysticerci) and Eimeria spp. were unrelated to sex, age or month; but Taenia intensity was highest among juveniles, and Eimeria intensity tended to decrease from December to April. Intensities of Nematodirus, Protostrongylus and Eimeria were higher in male hares than in females. Prevalence and intensity were correlated directly in Obeliscoides, Nematodirus, Trichuris and Eimeria. Hares that died during trapping and handling, or from natural predation, had greater intensities of Obeliscoides than did animals killed on purpose. There was no indication, however, that risk of death was increased by the other parasitic infections. Age-related immune responses to parasitism (except Obeliscoides) were evidenced by reduced or stabilized prevalence and/or intensity among older hares. A multiple-regression model predicted depressed body weight with increasing intensities of Nematodirus, Trichuris or Protostrongylus. Other body-condition and reproductive indices were unassociated with parasite intensities. Within the hare population, Obeliscoides, Trichuris, Protostrongylus and Taenia had overdispersed distributions (typical of many endoparasites) that did not differ from a negative binomial. The frequency with which each possible combination of helminth species occurred within individual hares was consistent with the assumption that such infections occurred independently. There was no compelling reason to believe parasitism was a significant factor in the overwinter decline of this population of snowshoe hares.     

Molecular Approaches Revealing Prehistoric, Historic, or Recent Translocations and Introductions of Hares (Genus Lepus) by Humans

Although only of medium size, and thus of little nutritional value compared to big game such as mammoths and ungulates, hares (Lepus spp.) probably have always been a food source for humans, as documented in archaeological finds. Nowadays, hares, particularly such species as the brown hare (L. europaeus), are among the most important game species in many European countries. For hunting, perhaps religious reasons, and in connection with certain myths, hares have been and are still being intentionally translocated. Ancient translocations by humans can be inferred from the presence of hares on islands that had no mainland connections, at least during the Pleistocene, the major evolutionary period of the genus Lepus. We review some of the literature on anthropogenic translocations of hares. We focus on three examples [the brown hare (L. europaeus), the Corsican hare (L. corsicanus), and the Sardinian hare (L. capensis)], where some molecular data could be used to trace the translocation routes and possible origins of introduced hare populations. Certain molecular marker systems, such as sequences of the hypervariable part I (HV-1) of the mitochondrial control region, show high variability in hare species and are thus promising for tracing both recent and ancient origins of translocated hares. Some other molecular marker systems as well as caveats connected with the use of such marker systems in the genus Lepus are also discussed.     

Helminth communities of an introduced hare (Lepus granatensis) and a native hare (Lepus europaeus) in southern France

We investigated the parasite communities of introduced Iberian hares (Lepus granatensis) and native European hares (Lepus europaeus) in southern France, where Iberian hares were introduced locally 20 yr ago as a game animal. Parasite communities of sympatric populations of the two hare species and of allopatric populations of European hares were compared. Iberian hares in France harbored a depauperate community of parasites relative to the population in its native habitat in Spain. European hares in areas of sympatry also were infected by Nematodiroides zembrae, which normally infects Iberian hares on their native range.     

The effect of supplemental food on the growth rates of neonatal,young, and adult cotton rats (Sigmodon hispidus) in northeastern Kansas,USA

In food-limited populations, the presence of extra food resources can influence the way individuals allocate energy to growth and reproduction. We experimentally increased food available to cotton rats (Sigmodon hispidus) near the northern limit of their range over a 2-year period and tested the hypothesis that seasonal growth rates would be enhanced by supplemental food during winter and spring when natural food levels are low. We also examined whether additional food resources were allocated to somatic growth or reproductive effort by pregnant and lactating females. The effect of supplemental food on growth varied with mass and season, but did not influence the growth rates of most cotton rats during spring and winter. In winter, small animals on supplemented grids had higher growth rates than small animals on control grids, but females in spring had lower growth rates under supplemented conditions. Growth rates of supplemented cotton rats were enhanced in summer. Northern cotton rat populations may use season-specific foraging strategies, maximizing energy intake during the reproductive season and minimizing time spent foraging in winter. Adult females invest extra resources in reproduction rather than in somatic growth. Pregnant females receiving supplemental food had higher growth rates than control females, and dependent pups (≤ 1 month of age) born to supplemented mothers had higher growth rates than those born to control mothers. Increased body size seems to confer an advantage during the reproductive season, but has no concomitant advantage to overwinter survival.