Arctic Ungulates Research Articles

Direct and indirect costs of parasitism preceding a population decline of an Arctic ungulate

Parasites negatively affect the fitness of ungulate hosts directly, and in wild ungulates, these effects may be synzootic with other stressors, such as limited nutritional resources. In the Arctic, muskoxen (Ovibos moschatus) occur in a highly seasonal environment and must rely on finite energetic resources for survival and productivity. We investigated the costs of gastrointestinal nematodes on the body condition and reproductive status of 141 muskoxen, on Banks Island, Canada, when the population was at a peak in numbers and density. Using a Partial Least Squares Path Modelling approach, we found that high adult nematode abundance was associated with lower body condition, and high parasite abundance was associated with female reproduction including the indirect effect through on body condition (n = 87). These findings suggest that individuals prioritize energetic reserves for reproduction over parasite defence. In fall 2003, a severe icing event that restricted access to forage was associated with high overwinter mortality of muskoxen and a population crash. Through direct and indirect costs of parasite infection on body condition and reproduction, the high abundance of parasites may have contributed to the effects of this extreme weather event. Understanding the mechanisms in which parasites impact fitness can help explain the ecological drivers of ungulate populations and predict the interactions between the environment and populations.

Weak coupling between energetic status and the timing of reproduction in an Arctic ungulate

Bioenergetic constraints are the ultimate determinant of the timing of reproduction, and seasonal breeding is consequently a widely observed trait. Consistent with this, attention has focused on plasticity in reproductive phenology conceptualized as a response to concomitant advances in the phenology of the environmental energy supply caused by climate change. Few studies, however, have directly compared timing of reproduction with energetic status in free-living wild animals. Here we demonstrate that neither body mass nor adiposity are strong proximate predictors of date of conception in wild reindeer (Rangifer tarandus). Weak coupling between energetic status and the phenology of reproduction accounts for the increasing discrepancy between the phenology of forage (energy supply) and the phenology of reproduction (energy demand) observed across the last 2–4 decades in two populations of this species. The results emphasise that phenological plasticity is not a passive response to changes in energy supply but derives from the way in which environmental factors interact with the core control mechanisms that govern timing. Central in this respect is integration, within the rheostatic centres of the hypothalamus, of information on nutritional status with the circannual life-history calendar.

High seasonal overlap in habitat suitability in a non-migratory High Arctic ungulate

Understanding drivers of space use and habitat selection is essential for management and conservation, especially under rapid environmental change. Here, we develop summer and winter habitat suitability models for the endemic wild Svalbard reindeer (Rangifer tarandus platyrhynchus). The High Arctic Svalbard tundra is currently subject to the fastest temperature increases on Earth, and reindeer spatial responses to associated environmental change are strongly restricted due to landscape barriers (including 60% glacial coverage) and lack of sea ice as movement corridors. We used an extensive dataset of GPS-collared adult females (2009–2018; N = 268 individual-years) to model seasonal habitat selection as a function of remotely sensed environmental variables , and subsequently built habitat suitability models using an ensemble modelling framework. As expected, we found that reindeer preferred productive habitats, described by the normalized difference vegetation index (NDVI) and plant biomass (derived from a vegetation map), in both seasons. This was further supported by selection for bird cliff areas, rich in forage, improving habitat suitability especially in winter. Contrary to our expectations, the terrain variables had similar, impact on habitat suitability in the two seasons, except for use of higher elevations in winter, likely related to improved forage access due to less snow. Suitable habitat patches covered only a small proportion of the landscape and were highly clustered in both seasons. About 13.0% of the total land area was suitable in both seasons, while summer-only and winter-only areas contributed a marginal addition of around 4.7% and 1.5%, respectively. This suggests, that unlike many continental and migratory Rangifer populations, even small geographic areas may encompass suffiscient suitable habitat. These first archipelago-wide habitat suitability models provide seasonal baseline maps relevant for the management and conservation of Svalbard reindeer, particularly under rapid environmental alterations from climate change.

Functional connectivity of an imperilled Arctic ungulate – where melting sea ice and human trails increase isolation

Sea ice loss, disturbance, and habitat modification by humans can alter functional landscape connectivity, with negative impacts on wildlife. Connectivity facilitates movement and gene flow, and contributes to genetic diversity, metapopulation dynamics, and species range-shifts under climate change. For Arctic ungulates, which disperse over large areas including sea ice, environmental change threatens further isolation. Protecting habitat and its linkages is critical and depends on identifying such areas at commensurate, broad scales. Using caribou, the world's most vagile species, we modelled and mapped the drivers of connectivity across ca. 2 million km2 of the Canadian Arctic Archipelago — where the pace of climate change is among the fastest and caribou are threatened with extinction. First, we quantified hierarchical genetic structure and identified two discrete groups. Next, using circuit theory and simultaneous multi-surface optimization, we tested whether land- and sea-scape heterogeneity or geographic distance better accounted for movement and gene flow within each group. We show that anthropogenic interference is far-reaching. High Arctic Peary caribou displayed isolation-by-resistance, where glacier cover, low sea-ice concentrations during fall, and human trails impeded connectivity. In contrast, more southerly barren-ground caribou displayed largely unrestricted gene flow. These divergent outcomes underscore that organism-landscape relationships can vary across space and highlight the importance of intra-specific structure and responses. By leveraging genetic data, our study demonstrates how critical movement pathways can be identified, even for remote and imperilled species. Such knowledge supports broad-scale conservation planning, in particular, by accounting for complex organism-landscape relationships, across vast, heterogeneous ecosystems.

Dynamic selection for forage quality and quantity in response to phenology and insects in an Arctic ungulate.

Spatiotemporal variation in forage is a primary driver of ungulate behavior, yet little is known about the nutritional components they select, and how selection varies across the growing season with changes in forage quality and quantity. We addressed these uncertainties in barren‐ground caribou (Rangifer tarandus), which experience their most important foraging opportunities during the short Arctic summer. Recent declines in Arctic caribou populations have raised concerns about the influence of climate change on summer foraging opportunities, given shifting vegetation conditions and insect harassment, and their potential effects on caribou body condition and demography. We examined Arctic caribou selection of summer forage by pairing locations from females in the Central Arctic Herd of Alaska with spatiotemporal predictions of biomass, digestible nitrogen (DN), and digestible energy (DE). We then assessed selection for these nutritional components across the growing season at landscape and patch scales, and determined whether foraging opportunities were constrained by insect harassment. During early summer, at the landscape scale, caribou selected for intermediate biomass and high DN and DE, following expectations of the forage maturation hypothesis. At the patch scale, however, caribou selected for high values of all forage components, particularly DN, suggesting that protein may be limiting. During late summer, after DN declined below the threshold for protein gain, caribou exhibited a switch at both spatial scales, selecting for higher biomass, likely enabling mass and fat deposition. Mosquito activity strongly altered caribou selection of forage and increased their movement rates, while oestrid fly activity had little influence. Our results demonstrate that early and late summer periods afford Arctic caribou distinct foraging opportunities, as they prioritize quality earlier in the summer and quantity later. Climate change may further constrain caribou access to DN as earlier, warmer Arctic summers may be associated with reduced DN and increased mosquito harassment.

Don't go chasing the ghosts of the past: habitat selection and site fidelity during calving in an Arctic ungulate

Predator avoidance and food availability are both factors known to influence habitat selection and site fidelity around calving in caribou and reindeer. Here, we assess habitat selection and site fidelity during the calving period in the solitary, Arctic Svalbard reindeer Rangifer tarandus platyrhynchus, which is subject to limited predation risk and human disturbance. In this largely predator-free environment, we explore and discuss if habitat selection during the first week after calving is driven by food availability or remnants of anti-predatory behaviour. Based on GPS-collar data and ultrasound scanning (2009–2017; n = 134 individual-years) from two study areas, we estimated individual calving dates using recursive partitioning and first passage time and compared habitat selection and site fidelity of reproductive versus non-reproductive females. The K-select analysis suggested similar habitat selection during calving in reproductive and non-reproductive females. Female reindeer generally selected for lowland, flat habitats with high proportion of heath and moss tundra, i.e. habitats typically rich in terms of forage plants. Individuals producing a calf had significantly higher site fidelity in the calving period compared to the null model, and the mean distance between consecutive years' calving areas ranged between 1.5 and 3.9 km. Our study provides support for the prediction that in the absence of significant predation, ungulate calving site selection in the Arctic is mainly driven by the availability of spatially and temporally varying food resources.

The neglected season: Warmer autumns counteract harsher winters and promote population growth in Arctic reindeer.

Arctic ungulates are experiencing the most rapid climate warming on Earth. While concerns have been raised that more frequent icing events may cause die-offs, and earlier springs may generate a trophic mismatch in phenology, the effects of warming autumns have been largely neglected. We used 25years of individual-based data from a growing population of wild Svalbard reindeer, to test how warmer autumns enhance population growth. Delayed plant senescence had no effect, but a six-week delay in snow-onset (the observed data range) was estimated to increase late winter body mass by 10%. Because average late winter body mass explains 90% of the variation in population growth rates, such a delay in winter-onset would enable a population growth of r=0.20, sufficient to counteract all but the most extreme icing events. This study provides novel mechanistic insights into the consequences of climate change for Arctic herbivores, highlighting the positive impact of warming autumns on population viability, offsetting the impacts of harsher winters. Thus, the future for Arctic herbivores facing climate change may be brighter than the prevailing view.

Quantifying energetic and fitness consequences of seasonal heterothermy in an Arctic ungulate.

Animals have adapted behavioral and physiological strategies to conserve energy during periods of adverse conditions. Heterothermy is one such adaptation used by endotherms. While heterothermy—fluctuations in body temperature and metabolic rate—has been shown in large vertebrates, little is known of the costs and benefits of this strategy, both in terms of energy and in terms of fitness. Hence, our objective was to model the energetics of seasonal heterothermy in the largest Arctic ungulate, the muskox (Ovibos moschatus), using an individual‐based energy budget model of metabolic physiology. We found that the empirically based drop in body temperature (winter max ~−0.8°C) overwinter in adult females resulted in substantial fitness benefits in terms of reduced daily energy expenditure and body mass loss. Body mass and energy reserves were 8.98% and 14.46% greater in modeled heterotherms compared to normotherms by end of winter. Based on environmental simulations, we show that seasonal heterothermy can, to some extent, buffer the negative consequences of poor prewinter body condition or reduced winter food accessibility, leading to greater winter survival (+20%–30%) and spring energy reserves (+10%–30%), and thus increased probability of future reproductive success. These results indicate substantial adaptive short‐term benefits of seasonal heterothermy at the individual level, with potential implications for long‐term population dynamics in highly seasonal environments.

Behavioral, Physiological, Demographic and Ecological Impacts of Hematophagous and Endoparasitic Insects on an Arctic Ungulate.

Animals that deliver a toxic secretion through a wound or to the body surface without a wound are considered venomous and toxungenous, respectively. Hematophagous insects, such as mosquitoes (Aedes spp.), meet the criteria for venomous, and some endoparasitic insects, such as warble flies (Hypoderma tarandi), satisfy the definition for toxungenous. The impacts of these insects on their hosts are wide ranging. In the Arctic, their primary host is the most abundant ungulate, the caribou (Rangifer tarandus). The most conspicuous impacts of these insects on caribou are behavioral. Caribou increase their movements during peak insect harassment, evading and running away from these parasites. These behavioral responses scale up to physiological effects as caribou move to less productive habitats to reduce harassment which increases energetic costs due to locomotion, reduces nutrient intake due to less time spent foraging, and can lead to poorer physiological condition. Reduced physiological condition can lead to lower reproductive output and even higher mortality rates, with the potential to ultimately affect caribou demographics. Caribou affect all trophic levels in the Arctic and the processes that connect them, thus altering caribou demographics could impact the ecology of the region. Broadening the definitions of venomous and toxungenous animals to include hematophagous and endoparasitic insects should not only generate productive collaborations among toxinologists and parasitologists, but will also lead to a deeper understanding of the ecology of toxic secretions and their widespread influence.

Phenotypic plasticity following weaning: winter body mass trends and food intake in relation to temperature by an arctic ungulate

Our knowledge is limited on how weaned mammals adjust to environmental events in the absence of maternal assistance in the polar north. With the expectation that weaned offspring should express the adaptive potential of the mature phenotype, we determined their appetite and body mass responses to extreme cold. Between December and May when mean daily temperature (Ta, °C) was between − 48 and 2.3 we provided five naturally weaned reindeer calves (Rangifer tarandus L.) ad libitum a pelleted feed of 11% crude protein with access to snow only in individual fenced outdoor pens in interior Alaska (64° N 147° W). We predicted voluntary food intake would meet maintenance of body mass, but that extreme cold would result in an increase in appetite that would be compensated for by access to warm water. A linear mixed effect model indicated a moderate daily increase in body mass from 55.9 ± 1.6 kg on 9 December to 59.2 ± 0.6 kg on 14 May and a moderate increase in intake that was evidence for maintenance of thermal homeostasis. However, at both low (≤ 30) and high (≥ 10) Ta (°C) intake increased and we estimated a lower critical temperature of − 22 and upper critical temperature at − 10 that was indicative of high total-body insulation. Despite the appetite response at low Ta, provision of warm water did not affect intake. Thus, whereas adult reindeer restrict intake by mobilization of endogenous reserves the calf thermogenic response was an increase in appetite that indicated the importance of access to a food base for calves in winter.

On the interplay between hypothermia and reproduction in a high arctic ungulate

For free-ranging animals living in seasonal environments, hypometabolism (lowered metabolic rate) and hypothermia (lowered body temperature) can be effective physiological strategies to conserve energy when forage resources are low. To what extent such strategies are adopted by large mammals living under extreme conditions, as those encountered in the high Arctic, is largely unknown, especially for species where the gestation period overlaps with the period of lowest resource availability (i.e. winter). Here we investigated for the first time the level to which high arctic muskoxen (Ovibos moschatus) adopt hypothermia and tested the hypothesis that individual plasticity in the use of hypothermia depends on reproductive status. We measured core body temperature over most of the gestation period in both free-ranging muskox females in Greenland and captive female muskoxen in Alaska. We found divergent overwintering strategies according to reproductive status, where pregnant females maintained stable body temperatures during winter, while non-pregnant females exhibited a temporary decrease in their winter body temperature. These results show that muskox females use hypothermia during periods of resource scarcity, but also that the use of this strategy may be limited to non-reproducing females. Our findings suggest a trade-off between metabolically-driven energy conservation during winter and sustaining foetal growth, which may also apply to other large herbivores living in highly seasonal environments elsewhere.

Stabilizing selection and adaptive evolution in a combination of two traits in an arctic ungulate.

Stabilizing selection is thought to be common in wild populations and act as one of the main evolutionary mechanisms, which constrain phenotypic variation. When multiple traits interact to create a combined phenotype, correlational selection may be an important process driving adaptive evolution. Here, we report on phenotypic selection and evolutionary changes in two natal traits in a semidomestic population of reindeer (Rangifer tarandus) in northern Finland. The population has been closely monitored since 1969, and detailed data have been collected on individuals since they were born. Over the length of the study period (1969-2015), we found directional and stabilizing selection toward a combination of earlier birth date and heavier birth mass with an intermediate optimum along the major axis of the selection surface. In addition, we demonstrate significant changes in mean traits toward earlier birth date and heavier birth mass, with corresponding genetic changes in breeding values during the study period. Our results demonstrate evolutionary changes in a combination of two traits, which agree closely with estimated patterns of phenotypic selection. Knowledge of the selective surface for combinations of genetically correlated traits are vital to predict how population mean phenotypes and fitness are affected when environmentschange.

Keeping cool in the warming Arctic: thermoregulatory behaviour by Svalbard reindeer (Rangifer tarandus platyrhynchus)

In animals with long generation times, evolution of physiological and morphological traits may not be fast enough to keep up with rapid climate warming, but thermoregulatory behaviour can possibly serve as an important buffer mitigating warming effects. In this study, we investigated if the cold-adapted Svalbard reindeer (Rangifer tarandus platyrhynchus (Vrolik, 1829)) used cool bed sites as a thermoregulatory behaviour in the summer. We recorded habitat variables and ground temperature at 371 bed sites with random “control” sites 10 and 100 m distant. Using case-control logistic regression, we found that reindeer selected bed sites on cool substrates (snow and mire), as well as cold, dry ground on days with warm ambient temperatures, while they avoided such sites on cold days. Selection of both cool substrates and cool ground did not depend on age or sex. The study was conducted in an environment where neither predatory threat nor insect harassment influenced bed site selection. Our findings suggest that the thermal landscape is important for habitat selection of cold-adapted Arctic ungulates in summer. Thus, behavioural strategies may be important to mitigate effects of climate change, at least in the short term.

Already at the bottom? Demographic declines are unlikely further to undermine genetic diversity of a large Arctic ungulate: muskox, Ovibos moschatus (Artiodactyla: Bovidae)

AbstractLow genetic diversity is associated with low fitness and evolutionary potential, yet the demographic and life-history traits of some species contribute to low genetic diversity, without empirical evidence of negative impacts on fitness. Modelling past and future trajectories of genetic diversity under different demographic scenarios can provide insight into how genetic variation might impact population fitness. The muskox is an Arctic species that has undergone multiple population bottlenecks and, although populations have rebounded repeatedly, two large populations have recently declined by > 50%. It is unclear how these demographic patterns influence muskox genetic diversity and fitness. We compared the genetic diversity of Canadian muskox populations undergoing opposing population trends. Genotyping 84 mainland and 244 Victoria Island individuals at ten microsatellite loci revealed low genetic variation (Victoria Island, mean allelic richness 1.66, expected heterozygosity 0.16; mainland, mean allelic richness 2.58, expected heterozygosity 0.41), with no evidence of further reductions in diversity subsequent to recent demographic declines. Bayesian modelling showed that a 1900s bottleneck contributed to the lack of diversity in contemporary populations, and forward-in-time simulations suggested little effect on genetic diversity over the next 100 years. Muskoxen might have reached a genetic diversity minimum, and additional research will be needed to determine their capacity to adapt to rapid changes in selective pressures in a rapidly changing Arctic.

Using seasonal landscape models to predict space use and migratory patterns of an arctic ungulate

BackgroundCaribou in the Western Arctic Herd undertake one of the longest, remaining intact migrations of terrestrial mammals in the world. They are also the most important subsistence resource for many northern rural residents, who rely on the caribou’s migratory movements to bring them near for harvest. Migratory geography has never been static, but subsistence harvesters have reported recent shifts in migration away from areas where they traditionally occurred. The reasons behind these changes are not well-understood, but may be related to rapid climate change and anthropogenic disturbances.MethodsTo predict changes in distribution and shifting migratory areas over the past decade, we used GPS telemetry data from adult females to develop predictive ecological niche models of caribou across northwestern Alaska. We employed the machine-learning algorithm, TreeNet, to analyze interactive, multivariate relationships between telemetry locations and 37 spatial environmental layers and to predict the distributions of caribou during spring, calving season, insect-harassment season, late summer, fall, and winter from 2009 to 2017. Model results were analyzed to identify regions of repeated predicted use, quantify mean longitude, predict land cover selection, and track migratory changes over time.ResultsDistribution models accurately predicted caribou at a spatially-explicit, 500-m scale. Model analyses identified migratory areas that shifted annually across the region, but which predicted 4 main areas of repeated use. Niche models were defined largely by non-linear relationships with coastally-influenced, climatic variables, especially snow-free date, potential evapo-transpiration, growing season length, proximity to sea ice, winter precipitation and fall temperature. Proximity to roads and communities were also important and we predicted caribou to generally occur more than 20–100 km from these features.ConclusionsWestern Arctic Herd caribou were predicted to occur in warmer, snow-free and treeless areas that may provide conditions conducive for efficient travel and foraging. Rapidly changing seasonal climates and coastal influences that determine forage availability, and human impediments that slow or divert movements are related to geographically and phenologically dynamic migration patterns that may periodically shift caribou away from traditional harvest areas. An enhanced understanding of the geographic behavior of caribou over time could inform traditional harvests and help conserve important Western Arctic caribou migratory areas.

More frequent extreme climate events stabilize reindeer population dynamics

Extreme climate events often cause population crashes but are difficult to account for in population-dynamic studies. Especially in long-lived animals, density dependence and demography may induce lagged impacts of perturbations on population growth. In Arctic ungulates, extreme rain-on-snow and ice-locked pastures have led to severe population crashes, indicating that increasingly frequent rain-on-snow events could destabilize populations. Here, using empirically parameterized, stochastic population models for High-Arctic wild reindeer, we show that more frequent rain-on-snow events actually reduce extinction risk and stabilize population dynamics due to interactions with age structure and density dependence. Extreme rain-on-snow events mainly suppress vital rates of vulnerable ages at high population densities, resulting in a crash and a new population state with resilient ages and reduced population sensitivity to subsequent icy winters. Thus, observed responses to single extreme events are poor predictors of population dynamics and persistence because internal density-dependent feedbacks act as a buffer against more frequent events.

Temperature-dependent development and freezing survival of protostrongylid nematodes of Arctic ungulates: implications for transmission

BackgroundUmingmakstrongylus pallikuukensis and Varestrongylus eleguneniensis are two potentially pathogenic lungworms of caribou and muskoxen in the Canadian Arctic. These parasites are currently undergoing northward range expansion at differential rates. It is hypothesized that their invasion and spread to the Canadian Arctic Archipelago are in part driven by climate warming. However, very little is known regarding their physiological ecology, limiting our ability to parameterize ecological models to test these hypotheses and make meaningful predictions. In this study, the developmental parameters of V. eleguneniensis inside a gastropod intermediate host were determined and freezing survival of U. pallikuukensis and V. eleguneniensis were compared.MethodsSlug intermediate hosts, Deroceras laeve, were collected from their natural habitat and experimentally infected with first-stage larvae (L1) of V. eleguneniensis. Development of L1 to third-stage larvae (L3) in D. laeve was studied at constant temperature treatments from 8.5 to 24 °C. To determine freezing survival, freshly collected L1 of both parasite species were held in water at subzero temperatures from -10 to -80 °C, and the number of L1 surviving were counted at 2, 7, 30, 90 and 180 days.ResultsThe lower threshold temperature (T0) below which the larvae of V. eleguneniensis did not develop into L3 was 9.54 °C and the degree-days required for development (DD) was 171.25. Both U. pallikuukensis and V. eleguneniensis showed remarkable freeze tolerance: more than 80% of L1 survived across all temperatures and durations. Larval survival decreased with freezing duration but did not differ between the two species.ConclusionBoth U. pallikuukensis and V. eleguneniensis have high freezing survival that allows them to survive severe Arctic winters. The higher T0 and DD of V. eleguneniensis compared to U. pallikuukensis may contribute to the comparatively slower range expansion of the former. Our study advances knowledge of Arctic parasitology and provides ecological and physiological data that can be useful for parameterizing ecological models.

Advancing the match-mismatch framework for large herbivores in the Arctic: Evaluating the evidence for a trophic mismatch in caribou.

Climate-induced shifts in plant phenology may adversely affect animals that cannot or do not shift the timing of their reproductive cycle. The realized effect of potential trophic “mismatches” between a consumer and its food varies with the degree to which species rely on dietary income and stored capital. Large Arctic herbivores rely heavily on maternal capital to reproduce and give birth near the onset of the growing season but are they vulnerable to trophic mismatch? We evaluated the long-term changes in the temperatures and characteristics of the growing seasons (1970–2013), and compared growing conditions and dynamics of forage quality for caribou at peak parturition, peak lactation, and peak forage biomass, and plant senescence between two distinct time periods over 36 years (1977 and 2011–13). Despite advanced thaw dates (7−12 days earlier), increased growing season lengths (15−21 days longer), and consistent parturition dates, we found no decline in forage quality and therefore no evidence within this dataset for a trophic mismatch at peak parturition or peak lactation from 1977 to 2011–13. In Arctic ungulates that use stored capital for reproduction, reproductive demands are largely met by body stores deposited in the previous summer and autumn, which reduces potential adverse effects of any mismatch between food availability and timing of parturition. Climate-induced effects on forages growing in the summer and autumn ranges, however, do correspond with the demands of female caribou and their offspring to gain mass for the next reproductive cycle and winter. Therefore, we suggest the window of time to examine the match-mismatch framework in Arctic ungulates is not at parturition but in late summer-autumn, where the multiplier effects of small changes in forage quality are amplified by forage abundance, peak forage intake, and resultant mass gains in mother-offspring pairs.

Climate and density dependence cause changes in adult sex ratio in a large Arctic herbivore

AbstractVariation in adult sex ratio (ASR) affects population demography and dynamics of large mammals. The mechanisms behind this variation are largely unclear, but may be partly related to climatic drivers and density dependence operating differently on the adult male and female segments of the population. Here, we examine such drivers of annual changes in ASR in the predator‐free wild Svalbard reindeer (Rangifer tarandus platyrhynchus), a high Arctic subspecies whose population dynamics are shaped by climate. Using up to 35 year long time‐series of population count data from two populations, we disentangle drivers of fluctuations in ASR by first analyzing how climate and density dependence affect sex‐specific adult population growth rates through effects on mortality. There were a positive population size trend and an overall female bias in ASR throughout the study period. Increased winter precipitation, a proxy for the harshness of winter feeding conditions, was found to significantly reduce adult population growth rates through reduced survival in males, but not in females. However, increased population size tended to cause a stronger immediate decline in female than in male adult population growth rates. As a consequence, the female bias in ASR increased with harsher winter conditions and declined with higher population size. As expected from the increased frequency of rainy and icy winters due to climate warming, a recent trend toward increased female bias in ASR was evident. This demonstrates that climatic drivers of both short‐term fluctuations and long‐term trends in demography need to be accounted for in the management and population dynamic predictions of Arctic ungulates.

The influence of weather conditions during gestation on life histories in a wild Arctic ungulate.

The internal predictive adaptive response (internal PAR) hypothesis predicts that individuals born in poor conditions should start to reproduce earlier if they are likely to have reduced performance in later life. However, whether this is the case remains unexplored in wild populations. Here, we use longitudinal data from a long-term study of Svalbard reindeer to examine age-related changes in adult female life-history responses to environmental conditions experienced in utero as indexed by rain-on-snow (ROSutero). We show that females experiencing high ROSutero had reduced reproductive success only from 7 years of age, independent of early reproduction. These individuals were able to maintain the same annual reproductive success between 2 and 6 years as phenotypically superior conspecifics that experienced low ROSutero Young females born after high ROSutero engage in reproductive events at lower body mass (about 2.5 kg less) than those born after low ROSutero The mean fitness of females that experienced poor environmental conditions in early life was comparable with that of females exposed to good environmental conditions in early life. These results are consistent with the idea of internal PAR and suggest that the life-history responses to early-life conditions can buffer the delayed effects of weather on population dynamics.