An isotopic approach to measuring nitrogen balance in caribou

Nutritional restrictions in winter may reduce the availability of protein for reproduction and survival in northern ungulates. We refined a technique that uses recently voided excreta on snow to assess protein status in wild caribou (Rangifer tarandus) in late winter. Our study was the first application of this non-invasive, isotopic approach to assess protein status of wild caribou by determining dietary and endogenous contributions of nitrogen (N) to urinary urea. We used isotopic ratios of N (δ¹⁵N) in urine and fecal samples to estimate the proportion of urea N derived from body N (p-UN) in pregnant, adult females of the Chisana Herd, a small population that ranged across the Alaska-Yukon border. We took advantage of a predator-exclosure project to examine N status of penned caribou in April 2006. Lichens were the primary forage (>40%) consumed by caribou in the pen and δ¹⁵N of fiber tracked the major forages in their diets. The δ¹⁵N of urinary urea for females in the pen was depleted relative (−1.3 ± 1.0 parts per thousand [‰], ) to the δ¹⁵N of body N (2.7 ± 0.7‰). A similar proportion of animals in the exclosure lost core body mass (excluding estimates of fetal and uterine tissues; 55%) and body protein (estimated by isotope ratios; 54%). This non-invasive technique could be applied at various spatial and temporal scales to assess trends in protein status of free-ranging populations of northern ungulates. Intra- and inter-annual estimates of protein status could help managers monitor effects of foraging conditions on nutritional constraints in ungulates, increase the efficiency and efficacy of management actions, and help prepare stakeholders for potential changes in population trends. © 2010 The Wildlife Society.     

Calving rate,calf survival rate,and habitat selection of forest-dwelling caribou in a highly managed landscape

Logging negatively affects the threatened forest-dwelling caribou (Rangifer tarandus caribou) through its positive effects on large predator populations. As recruitment is a key component of caribou population growth rate, we assessed calving rates of females and calf survival rates during the most critical period for calf survival, the calving period. We also identified causes of calf mortality and investigated the influence of predation risk, food availability, and human disturbance on habitat selection of females during the calving period at both the home-range and forest stand scales. We hypothesized that caribou should display habitat selection patterns to reduce predation risk at both scales. Using telemetry, we followed 22 females and their calves from 2004 to 2007 in a highly managed study area in Québec, Canada. Most females (78.5 ± 0.05 [SE]) gave birth each year, but only 46.3 ± 8.0% of the calves survived during the first 50 days following birth, and 57.3 ± 14.9% of them died from black bear (Ursus americanus) predation. At the home-range scale, caribou selected calving areas located at upper slope positions and avoided high road density areas. Surprisingly, they also selected the forested habitat type having the lowest lateral cover (mixed and deciduous stands) while avoiding the highest cover (regenerating conifer stands). At the forest stand scale, caribou selected areas located at relatively high elevations and with a lower basal area of black spruce trees. The selection of upper slope positions likely favored spatial segregation between calving females and wolves (Canis lupus) but not black bear. Our results suggest that calving females used areas from which they could visually detect approaching predators. While wolf avoidance appeared to be effective in a highly managed landscape, caribou did not appear to have adjusted their predator avoidance strategy to the recent increase in black bear abundance, who have benefited from increased food abundance. This situation requires focused attention from wildlife managers as logging activities are progressing towards the north within the core of forest-dwelling caribou range. © 2011 The Wildlife Society.     

Comparison of Woodland Caribou Calving Areas Determined by Movement Patterns Across Northern Ontario

Adult female survival and calf recruitment influence population dynamics, but there is limited information on calving and neonatal mortality of boreal woodland caribou (Rangifer tarandus caribou; caribou) in Ontario, Canada. We identified calf parturition sites and 5-week neonatal mortality using a movement-based approach across 3 northern Ontario study regions (Pickle Lake, Nakina, and Cochrane) that vary in their capacity to support caribou populations. In comparing 22 caribou-years of video-collar footage during 2010–2013 to predictions of the movement-based approach, we found live parturition events were 100% correctly classified, date of parturition was within 1.08 ± 0.28 ( ± SE) days, and mortality events up to 5 weeks postpartum were 88% correctly classified. Across study regions, 87% of 186 caribou were pregnant and 76% of 107 caribou-years indicated birth events with median parturition dates a week later in Cochrane (23 May) than in Pickle Lake (17 May) and Nakina (16 May). Based on selection ratios of caribou-years with calves-at-heel (n = 80), caribou consistently selected for lowlands and closed-canopied forests and mostly against early-seral stands (<20 yrs old) and areas near linear features during the neonatal and the post-neonatal period (up to 35 days postpartum). Based on the video footage and movement models, 30% of 81 caribou-years that indicated live births also showed females lost their calf within the first 5 weeks postpartum, with higher risk of neonatal mortality associated with increased use of lowlands and greater postpartum movement rates. This study provides informative metrics of caribou reproduction across northern Ontario that will contribute to future population modeling and identifies important landscape features to be considered in future industrial development and land use planning for caribou conservation. © 2020 The Wildlife Society.     

Snow depth does not affect recruitment in a low-density population of boreal woodland caribou (<Emphasis Type="Italic">Rangifer tarandus caribou</Emphasis>)

Winter snow depth may be an important driver of annual variability in recruitment of ungulate calves, and low calf recruitment has been implicated as a factor in declining boreal caribou (Rangifer tarandus caribou) populations. We used 11 consecutive years (2006–2016) of aerial survey data to document calf recruitment in a low-density population of boreal woodland caribou in the Northwest Territories, Canada. We measured snow depth in winter and tested two hypotheses: (1) that calf recruitment was lower in winters with greater snow depth and (2) that calf recruitment was lower following winters with greater snow depth (1-year time lag). Recruitment, the number of calves/adult female in March, ranged twofold from 0.23 to 0.45, and snow depth also ranged twofold from 41 to 85 cm. Yet, we found no support for the hypothesis that late-winter snow depth in the current or previous year was inversely related to calf recruitment.     

Divergent estimates of herd‐wide caribou calf survival: Ecological factors and methodological biases

Population monitoring is a critical part of effective wildlife management, but methods are prone to biases that can hinder our ability to accurately track changes in populations through time. Calf survival plays an important role in ungulate population dynamics and can be monitored using telemetry and herd composition surveys. These methods, however, are susceptible to unrepresentative sampling and violations of the assumption of equal detectability, respectively. Here, we capitalized on 55 herd‐wide estimates of woodland caribou (Rangifer tarandus caribou) calf survival in Newfoundland, Canada, using telemetry (n = 1,175 calves) and 249 herd‐wide estimates of calf:cow ratios (C:C) using herd composition surveys to investigate these potential biases. These data included 17 herd‐wide estimates replicated from both methods concurrently (n = 448 calves and n = 17 surveys) which we used to understand which processes and sampling biases contributed to disagreement between estimates of herd‐wide calf survival. We used Cox proportional hazards models to determine whether estimates of calf mortality risk were biased by the date a calf was collared. We also used linear mixed‐effects models to determine whether estimates of C:C ratios were biased by survey date and herd size. We found that calves collared later in the calving season had a higher mortality risk and that C:C tended to be higher for surveys conducted later in the autumn. When we used these relationships to modify estimates of herd‐wide calf survival derived from telemetry and herd composition surveys concurrently, we found that formerly disparate estimates of woodland caribou calf survival now overlapped (within a 95% confidence interval) in a majority of cases. Our case study highlights the potential of under‐appreciated biases to impact our understanding of population dynamics and suggests ways that managers can limit the influence of these biases in the two widely applied methods for estimating herd‐wide survival.     

Relative costs of offspring sex and offspring survival in a polygynous mammal

Costs of reproduction are expected to be ubiquitous in wild animal populations and understanding the drivers of variation in these costs is an important aspect of life-history evolution theory. We use a 43 year dataset from a wild population of red deer to examine the relative importance of two factors that influence the costs of reproduction to mothers, and to test whether these costs vary with changing ecological conditions. Like previous studies, our analyses indicate fitness costs of lactation: mothers whose calves survived the summer subsequently showed lower survival and fecundity than those whose calves died soon after birth, accounting for 5% and 14% of the variation in mothers'' survival and fecundity, respectively. The production of a male calf depressed maternal survival and fecundity more than production of a female, but accounted for less than 1% of the variation in either fitness component. There was no evidence for any change in the effect of calf survival or sex with increasing population density.     

Avoidance of roads and selection for recent cutovers by threatened caribou: fitness-rewarding or maladaptive behaviour?

The impact of anthropogenic disturbance on the fitness of prey should depend on the relative effect of human activities on different trophic levels. This verification remains rare, however, especially for large animals. We investigated the functional link between habitat selection of female caribou (Rangifer tarandus) and the survival of their calves, a fitness correlate. This top-down controlled population of the threatened forest-dwelling caribou inhabits a managed forest occupied by wolves (Canis lupus) and black bears (Ursus americanus). Sixty-one per cent of calves died from bear predation within two months following their birth. Variation in habitat selection tactics among mothers resulted in different mortality risks for their calves. When calves occupied areas with few deciduous trees, they were more likely to die from predation if the local road density was high. Although caribou are typically associated with pristine forests, females selected recent cutovers without negative impact on calf survival. This selection became detrimental, however, as regeneration took place in harvested stands owing to increased bear predation. We demonstrate that human disturbance has asymmetrical consequences on the trophic levels of a food web involving multiple large mammals, which resulted in habitat selection tactics with a greater short-term fitness payoff and, therefore, with higher evolutionary opportunity.     

Woodland caribou calf mortality in Newfoundland: insights into the role of climate,predation and population density over three decades of study

The rates and causes of juvenile mortality are central features of the dynamics and conservation of large mammals, like woodland caribou (Rangifer tarandus caribou (Gmelin, 1788)), but intrinsic and extrinsic factors may be modified by variations in animal abundance. We tested the influences of population size, climate, calf weight and sex on survival to 6 months of age of 1241 radio-collared caribou calves over three decades, spanning periods of population growth (1979–1997) and decline (2003–2012) in Newfoundland, Canada. Daily survival rates were higher and rose more quickly with calf age during the population growth period compared to the decline. Population size (negatively) and calf weight (positively) affected survival during the decline but neither had a detectable influence during the growth phase. Sex, climate and plant productivity (the latter two derived from the North Atlantic Oscillation and Normalized Difference Vegetation Index, respectively) exerted minimal influence during either phase. Predation was the dominant source of mortality. The mean percentage of calves killed by predators was 30 % higher during the decline compared to the growth phase. Black bears (Ursus americanus) and lynx (Lynx canadensis) were the major predators during the population increase but this changed during the decrease to black bears and coyotes (Canis latrans). Our findings are consistent with the hypothesis that Newfoundland caribou experienced phase-dependent survival mediated proximally by predation and competition for food.     

Understanding predation risk and individual variation in risk avoidance for threatened boreal caribou

Predation risk is a driver of species’ distributions. Animals can increase risk avoidance in response to fluctuations in predation risk, but questions remain regarding individual variability and the capacity to respond to changes in spatial risk across human‐altered landscapes. In northeast British Columbia, Canada, boreal caribou populations declined as roads and seismic lines have increased, which are theorized to increase gray wolf predation. Our goal was to model risk and to evaluate individual variability and the development of risk perception by examining individual risk avoidance in response to reproductive status and age. We used locations from collared caribou and wolves to identify landscape features associated with the risk of a potential wolf‐caribou encounter and risk of being killed given an encounter. We built resource selection functions to estimate individual responses to risk. We used general linear regressions to evaluate individual risk and linear feature avoidance as a function of age and reproductive status (calf or no calf). Linear features increased the risk of encounter. Older caribou and caribou with calves demonstrated stronger avoidance of the risk of encounter and roads, but weaker avoidance in late summer to the risk of being killed relative to younger and calf‐less individuals. Mechanisms explaining the inverse relationships between the risk of encounter and risk of being killed are uncertain, but it is conceivable that caribou learn to avoid the risk of encounter and roads. Responses by females with vulnerable calves to the risk of encounter and risk of being killed might be explained by a trade‐off between these two risk types and a prioritization on the risk of encounter. Despite the capacity to alter their responses to risk, the global decline in Rangifer populations (caribou and wild reindeer) suggests these behaviors are insufficient to mitigate the impacts of anthropogenic disturbances.     

Behavioural strategies towards human disturbances explain individual performance in woodland caribou

Behavioural strategies may have important fitness, ecological and evolutionary consequences. In woodland caribou, human disturbances are associated with higher predation risk. Between 2004 and 2011, we investigated if habitat selection strategies of female caribou towards disturbances influenced their calf’s survival in managed boreal forest with varying intensities of human disturbances. Calf survival was 53 % and 43 % after 30 and 90 days following birth, respectively, and 52 % of calves that died were killed by black bear. The probability that a female lose its calf to predation was not influenced by habitat composition of her annual home range, but decreased with an increase in proportion of open lichen woodland within her calving home range. At the local scale, females that did not lose their calf displayed stronger avoidance of high road density areas than females that lost their calf to predation. Further, females that lost their calf to predation and that had a low proportion of ≤5-year-old cutovers within their calving home range were mostly observed in areas where these young cutovers were locally absent. Also, females that lost their calf to predation and that had a high proportion of ≤5-year-old cutovers within their calving home range were mostly observed in areas with a high local density of ≤5-year-old cutovers. Our study demonstrates that we have to account for human-induced disturbances at both local and regional scales in order to further enhance effective caribou management plans. We demonstrate that disturbances not only impact spatial distribution of individuals, but also their reproductive success.     

Migratory herds of wildebeests and zebras indirectly affect calf survival of giraffes

相似文献   

Anthropogenic Disturbance and Population Viability of Woodland Caribou in Ontario

One of the most challenging tasks in wildlife conservation and management is to clarify how spatial variation in land cover due to anthropogenic disturbance influences wildlife demography and long-term viability. To evaluate this, we compared rates of survival and population growth by woodland caribou (Rangifer tarandus caribou) from 2 study sites in northern Ontario, Canada that differed in the degree of anthropogenic disturbance because of commercial logging and road development, resulting in differences in predation risk due to gray wolves (Canis lupus). We used an individual-based model for population viability analysis (PVA) that incorporated adaptive patterns of caribou movement in relation to predation risk and food availability to predict stochastic variation in rates of caribou survival. Field estimates of annual survival rates for adult female caribou in the unlogged ( 0.90) and logged ( 0.76) study sites recorded during 2010–2014 did not differ significantly (P > 0.05) from values predicted by the individual-based PVA model (unlogged: = 0.87; logged: 0.79). Outcomes from the individual-based PVA model and a simpler stage-structured matrix model suggest that substantial differences in adult survival largely due to wolf predation are likely to lead to long-term decline of woodland caribou in the commercially logged landscape, whereas the unlogged landscape should be considerably more capable of sustaining caribou. Estimates of population growth rates (λ) for the 2010–2014 period differed little between the matrix model and the individual-based PVA model for the unlogged (matrix model = 1.01; individual-based model = 0.98) and logged landscape (matrix model = 0.88; individual-based model = 0.89). We applied the spatially explicit PVA model to assess the viability of woodland caribou across 14 woodland caribou ranges in Ontario. Outcomes of these simulations suggest that woodland caribou ranges that have experienced significant levels of commercial forestry activities in the past had annual growth rates <0.89, whereas caribou ranges that had not experienced commercial forestry operations had population growth rates >0.96. These differences were strongly related to regional variation in wolf densities. Our results suggest that increased wolf predation risk due to anthropogenic disturbance is of sufficient magnitude to cause appreciable risk of population decline in woodland caribou in Ontario. © 2020 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

EXTINCTION-RECOLONIZATION DYNAMICS IN THE MYCOPHAGOUS BEETLE PHALACRUS SUBSTRIATUS

The population structure of the mycophagous beetle Phalacrus substriatus is characterized by many small, local populations interconnected by migration over a small spatial scale (10 × 75 m²). Each local P. substriatus population has a relatively short expected persistence time, but persistence of the species occurs due to a balance between frequent local extinctions and recolonizations. This nonequilibrium population structure can have profound effects on how the genetic variation is structured between and within populations. Theoretical models have stated that the genetic differentiation among local populations will be enhanced relative to an island model at equilibrium if the number of colonizers is less than approximately twice the number of migrants among local populations. To study these effects, a set of 50 local P. substriatus populations were surveyed over a four-year period to record any naturally occurring extinctions and recolonizations. The per population colonization and extinction rate were 0.237 and 0.275, respectively. Mark-recapture techniques were used to estimate a number of demographic parameters: local population size (N = 11.1), migration rate , number of colonizers (k = 4.0), and the probability of common origin of colonizers (φ = 0.5). The theoretically predicted level of differentiation among local populations (measured as Wright's F_ST) was 0.070. Genetic data obtained from an electrophoretic survey of seven polymorphic loci gave an estimated degree of differentiation of 0.077. There was thus a good agreement between the empirical results and the theoretical predictions. Young populations had significantly higher levels of differentiation than old, more established populations . The extinction-recolonization dynamics resulted in an overall increase in the genetic differentiation among local populations by c. 40%. The global effective population size was also reduced by c. 55%. The results give clear evidence to how nonequilibrium processes shape the genetic structure of populations.     

Partial genotyping at polymorphic markers can improve heritability estimates in sibling groups

Accurate estimates of heritability () are necessary to assess adaptive responses of populations and evolution of fitness‐related traits in changing environments. For plants, estimates generally rely on maternal progeny designs, assuming that offspring are either half‐sibs or unrelated. However, plant mating systems often depart from half‐sib assumptions, this can bias estimates. Here, we investigate how to accurately estimate in nonmodel species through the analysis of sibling designs with a moderate genotyping effort. We performed simulations to investigate how microsatellite marker information available for only a subset of offspring can improve estimates based on maternal progeny designs in the presence of nonrandom mating, inbreeding in the parental population or maternal effects. We compared the basic family method, considering or not adjustments based on average relatedness coefficients, and methods based on the animal model. The animal model was used with average relatedness information, or with hybrid relatedness information: associating one‐generation pedigree and family assumptions, or associating one‐generation pedigree and average relatedness coefficients. Our results highlighted that methods using marker‐based relatedness coefficients performed as well as pedigree‐based methods in the presence of nonrandom mating (i.e. unequal male reproductive contributions, selfing), offering promising prospects to investigate in situ heritabilities in natural populations. In the presence of maternal effects, only the use of pairwise relatednesses through pedigree information improved the accuracy of estimates. In that case, the amount of father‐related offspring in the sibling design is the most critical. Overall, we showed that the method using both one‐generation pedigree and average relatedness coefficients was the most robust to various ecological scenarios.     

Impact of Spatial and Temporal Variation in Calf Survival on the Growth of Elk Populations

Abstract: The realized impact of a vital rate on population growth (λ) is determined by both the relative influence of the vital rate on λ (elasticity) and its magnitude of variability. We estimated mean survival and reproductive rates in elk (Cervus elaphus) and spatial and temporal variation in these rates from 37 sources located primarily across the Rocky Mountain region and northwestern United States. We removed sampling variance from estimates of process variance both within and across vital-rate data sets using the variance discounting method developed by White (2000). Deterministic elasticities calculated from a population matrix model parameterized with these mean vital rates ranked adult female survival (e_Scow = 0.869) much higher than calf survival (e_Scalf = 0.131). However, process variance in calf survival was >11 times greater than process variance in female survival across data sets and 10 times greater on average within studies. We conducted Life-Stage Simulation Analysis to incorporate both vital-rate elasticity patterns and empirical estimates of variability to identify those vital rates most influential in elk population dynamics. The overwhelming magnitude of variation in calf survival explained 75% of the variation in the population growth rates generated from 1,000 matrix replicates, compared to just 16% of the variation in λ explained by variation in female survival. Variation in calf survival greatly impacts elk population growth and calls into question the utility of classical elasticity analysis alone for guiding elk management. These results also suggest that the majority of interannual variability that wildlife managers document in late-winter and spring elk surveys is attributable to variation in calf survival over the previous year and less influenced by variation in the harvest of females during the preceding autumn. To meet elk population size objectives, managers should consider the inherent variation in calf survival, and its apparent sensitivity to management, in addition to female harvest.     

Reproduction,birth seasonality,and calf survival of bottlenose dolphins in Doubtful Sound,New Zealand

Understanding the reproductive parameters of very small or declining populations is of clear importance to conservation. From 1995 to 2011 we recorded calf production (n = 71) and calf survival for 27 breeding females in the bottlenose dolphin (Tursiops truncatus) population in Doubtful Sound, New Zealand; a population with a recent history of declining abundance. Overall, 67% of calves survived their first year, and 40% survived to 3 yr (or are 2 yr old and still alive). Most calves that died in the first year died in their first month (87%). Multiparous mothers (n = 18) showed high variation in calf survival. The most successful six had all but one of their 20 calves (95%) survive to 1 yr. Fourteen of the 20 (70%) survived to 3 yr, and another four are still alive and are 1 or 2 yr old. In contrast, the least successful seven mothers produced a similar number of calves (21), eight of which (38%) survived to 1 yr, and none to 3 yr. Here we describe calving seasonality and calf survival, observed over 16 yr, and show that large variation in reproductive success of individual females is an example of extreme demographic stochasticity in this small, endangered population.     

Group characteristics,site fidelity,and photo‐identification of harbor porpoises,Phocoena phocoena,in Burrows Pass,Fidalgo Island,Washington

Little is known about harbor porpoises at the individual level or local group structure. Group characteristics, site fidelity, and photo‐identification of harbor porpoises were investigated off Fidalgo Island, Washington State. Harbor porpoise presence was affected by season and rip tide strength (Wald χ² P < 0.04); calf presence was influenced by season and tide (Wald χ² P < 0.0075). Average group size (2.32 ± 1.38, n = 266) was influenced by season, behavior, and calf presence (F₇ = 9.71, P < 0.0001, R² = 0.294). Fifty‐three individuals were identified using a matrix of primary, secondary, and confirmation markings that were stable over months/years. Over 35% were resighted in more than 1 mo (range 1–7, = 1.83); 15.1% were seen in more than 1 yr, suggesting some level of residency. Despite having higher effort, presence and group size were significantly lower in Summer. Variations in the significance of rip tide strength and tides relate to calf presence and support other findings that harbor porpoise population structure is complex and varies at small spatiotemporal scales and may also vary between populations and habitats. This study identifies variables affecting group characteristics and emphasizes the importance of research on local populations of harbor porpoises.     

Woodland caribou habitat selection patterns in relation to predation risk and forage abundance depend on reproductive state

The ideal free distribution assumes that animals select habitats that are beneficial to their fitness. When the needs of dependent offspring differ from those of the parent, ideal habitat selection patterns could vary with the presence or absence of offspring. We test whether habitat selection depends on reproductive state due to top‐down or bottom‐up influences on the fitness of woodland caribou (Rangifer tarandus caribou), a threatened, wide‐ranging herbivore. We combined established methods of fitting resource and step selection functions derived from locations of collared animals in Ontario with newer techniques, including identifying calf status from video collar footage and seasonal habitat selection analysis through latent selection difference functions. We found that females with calves avoided predation risk and proximity to roads more strongly than females without calves within their seasonal ranges. At the local scale, females with calves avoided predation more strongly than females without calves. Females with calves increased predation avoidance but not selection for food availability upon calving, whereas females without calves increased selection for food availability across the same season. These behavioral responses suggest that habitat selection by woodland caribou is influenced by reproductive state, such that females with calves at heel use habitat selection to offset the increased vulnerability of their offspring to predation risk.     

Effective size of density‐dependent populations in fluctuating environments

Reliable estimates of effective population size are of central importance in population genetics and evolutionary biology. For populations that fluctuate in size, harmonic mean population size is commonly used as a proxy for (multi‐) generational effective size. This assumes no effects of density dependence on the ratio between effective and actual population size, which limits its potential application. Here, we introduce density dependence on vital rates in a demographic model of variance effective size. We derive an expression for the ratio in a density‐regulated population in a fluctuating environment. We show by simulations that yearly genetic drift is accurately predicted by our model, and not proportional to as assumed by the harmonic mean model, where N is the total population size of mature individuals. We find a negative relationship between and N. For a given N, the ratio depends on variance in reproductive success and the degree of resource limitation acting on the population growth rate. Finally, our model indicate that environmental stochasticity may affect not only through fluctuations in N, but also for a given N at a given time. Our results show that estimates of effective population size must include effects of density dependence and environmental stochasticity.     

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.