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1.The cost of reproduction on demographic rates is often assumed to operate through changing body condition. Several studies have found that reproduction depresses body mass more if the current conditions are severe, such as high population densities or adverse weather, than under benign environmental conditions. However, few studies have investigated the association between the fitness components and body mass costs of reproduction. 2.Using 25 years of individual-based capture–recapture data from Svalbard reindeer Rangifer tarandus platyrhynchus, we built a novel Bayesian state-space model that jointly estimated interannual change in mass, annual reproductive success and survival, while accounting for incomplete observations. The model allowed us to partition the differential effects of intrinsic and extrinsic factors on both non-reproductive mass change and the body mass cost of reproduction, and to quantify their consequences on demographic rates.3. Contrary to our expectation, the body mass cost of reproduction (mean = –5.8 kg) varied little between years (CV = 0.08), whereas the between-year variation in body mass changes, that were independent of the previous year's reproductive state, varied substantially (CV = 0.4) in relation to autumn temperature and the amount of rain-on-snow in winter. This body mass loss led to a cost of reproduction on the next reproduction, which was amplified by the same environmental covariates, from a 10% reduction in reproductive success in benign years, to a 50% reduction in harsh years. The reproductive mass loss also resulted in a small reduction in survival.4. Our results show how demographic costs of reproduction, driven by interannual fluctuations in individual body condition, result from the balance between body mass costs of reproduction and body mass changes that are independent of previous reproductive state. We illustrate how a strong context-dependent fitness cost of reproduction can occur, despite a relatively fixed body mass cost of reproduction. This suggests that female reindeer display a very conservative energy allocation strategy, either aborting their reproductive attempt at an early stage or weaning at a relatively constant cost. Such a strategy might be common in species living in a highly stochastic and food limited environment.

Significance Whether the renowned population cycles of small mammals in northern food webs are driven by bottom-up (plant–herbivore) or top-down (predator–prey) interactions is still a debated question but crucial to our understanding of their ecological functions and response to climate change. A long-term study of a graminivorous vole population in an exceptionally simple High Arctic food web allowed us to identify which population dynamics features are present without top-down regulation. Unique features were high-amplitude, noncyclic population fluctuations driven by a combination of stochastic weather events and season-specific density dependence likely arising from plant–herbivore interactions. That such features are not present in more complex food webs points to the importance of top-down regulation in small mammal populations., Ecologists are still puzzled by the diverse population dynamics of herbivorous small mammals that range from high-amplitude, multiannual cycles to stable dynamics. Theory predicts that this diversity results from combinations of climatic seasonality, weather stochasticity, and density-dependent food web interactions. The almost ubiquitous 3- to 5-y cycles in boreal and arctic climates may theoretically result from bottom-up (plant–herbivore) and top-down (predator–prey) interactions. Assessing, empirically, the roles of such interactions and how they are influenced by environmental stochasticity has been hampered by food web complexity. Here, we take advantage of a uniquely simple High Arctic food web, which allowed us to analyze the dynamics of a graminivorous vole population not subjected to top-down regulation. This population exhibited high-amplitude, noncyclic fluctuations—partly driven by weather stochasticity. However, the predominant driver of the dynamics was overcompensatory density dependence in winter that caused the population to frequently crash. Model simulations showed that the seasonal pattern of density dependence would yield regular 2-y cycles in the absence of stochasticity. While such short cycles have not yet been observed in mammals, they are theoretically plausible if graminivorous vole populations are deterministically bottom-up regulated. When incorporating weather stochasticity in the model simulations, cyclicity became disrupted and the amplitude was increased—akin to the observed dynamics. Our findings contrast with the 3- to 5-y population cycles that are typical of graminivorous small mammals in more complex food webs, suggesting that top-down regulation is normally an important component of such dynamics.

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. bed site, behavioural buffering, climate change, habitat selection, snow, ungulates, Svalbard reindeer, Rangifer tarandus platyrhynchus.

Ecologists are still puzzled by the diverse population dynamics of herbivorous small mammals that range from high-amplitude, multi-annual cycles to stable dynamics. Theory predicts that this diversity results from combinations of climatic seasonality, weather stochasticity and density-dependent food web interactions. The almost ubiquitous 3-5-yr cycles in boreal and arctic climates may theoretically result from bottom-up (plant-herbivore) and top-down (predator-prey) interactions. Assessing empirically the roles of such interactions, and how they are influenced by environmental stochasticity, has been hampered by food web complexity. Here, we take advantage of a uniquely simple High-Arctic food web, which allowed us to analyze dynamics of a graminivorous vole population not subjected to top-down regulation. This population exhibited high-amplitude, non-cyclic fluctuations - partly driven by weather stochasticity. However, the predominant driver of the dynamics was direct density dependence, which alternated between being weak in summer and strong (overcompensatory) in winter that the population frequently crashed. Model simulations showed that this season-specific density dependence would yield regular 2-year cycles in absence of stochasticity. While such short cycles have not yet been observed in mammals, they are theoretically plausible if graminivorous vole populations are deterministically bottom-up regulated. When incorporating weather stochasticity in the model simulations, cyclicity became disrupted and the amplitude was increased - akin to the observed dynamics. Our findings contrast with the 3-5-yr population cycles involving delayed density dependence that are typical of graminivorous small mammals in more complex food webs, suggesting that top-down regulation is an important component of such dynamics.

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.

Seasonal energetic challenges may constrain an animal's ability to respond to changing individual and environmental conditions. Here, we investigated variation in heart rate, a well-established proxy for metabolic rate, in Svalbard reindeer (Rangifer tarandus platyrhynchus), a species with strong seasonal changes in foraging and metabolic activity. In 19 adult females, we recorded heart rate, subcutaneous temperature and activity using biologgers. Mean heart rate more than doubled from winter to summer. Typical drivers of energy expenditure, such as reproduction and activity, explained a relatively limited amount of variation (2–6% in winter and 16–24% in summer) compared to seasonality, which explained 75% of annual variation in heart rate. The relationship between heart rate and subcutaneous temperature depended on individual state via body mass, age and reproductive status, and the results suggested that peripheral heterothermy is an important pathway of energy management in both winter and summer. While the seasonal plasticity in energetics makes Svalbard reindeer well-adapted to their highly seasonal environment, intraseasonal constraints on modulation of their heart rate may limit their ability to respond to severe environmental change. This study emphasizes the importance of encompassing individual state and seasonal context when studying energetics in free-living animals.This article is part of the theme issue ‘Measuring physiology in free-living animals (Part II)’.

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 25 years 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. body mass, climate change, fitness, GPS, movement ecology, plant phenology, Rangifer, snow, space use, ungulates © 2020 The Authors. Global Change Biology published by John Wiley & Sons Ltd This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

Question: Vascular plant productivity of arctic tundra has often been viewed as varying little between years and thus being largely insensitive to the high inter-annual variation in summer weather conditions. Yet, remote-sensing data and retrospective growth analyses of the circumpolar dwarf shrub, Cassiope tetragona, commonly show considerable between-year variability in plant growth in response to variation in summer temperature. Given that both Cassiope growth and vascular plant biomass production share a common environmental driver, summer temperature, we would expect positive co-variation between them. Here we investigate whether this is indeed the case and if so over what spatial scale. Location: Nordenskiöldland, high arctic Svalbard. Methods: We brought dendroecology and plot-based field estimation methodologies together in an empirical study using retrospective analysis of Cassiope growth and annual estimation of above-ground vegetation biomass production to investigate their temporal and spatial co-variation and sensitivity to summer weather conditions. Results: Despite substantial small-scale heterogeneity, we found spatial co-variation in Cassiope growth patterns, which weakened as distance between sampling sites increased from 0 to 25 km. Furthermore, we found a strong positive correlation between annual estimates of above-ground live vascular plant biomass and Cassiope shoot growth over a 12-year period at a local scale. The correlation declined with distance, likely due to increasing differences in local weather conditions. Conclusions: We demonstrate that Cassiope growth can be used as a proxy for above-ground tundra vegetation productivity at the local scale. Our findings suggest that Arctic plant productivity is as sensitive to between-year variation in summer temperature as the well-established growth response of Cassiope. This challenges the view that tundra plant productivity varies little between years and provides a mechanistic understanding that helps reconcile field-and satellite-based annual estimation methods. K E Y W O R D S Arctic bell-heather, arctic climate change, Cassiope tetragona, dendrochronology, plant– climate interaction, spatial scale, Svalbard, tundra vegetation, vascular plant productivity, vegetation biomass

Warming of the Arctic has resulted in earlier snowmelt and green-up of plants in spring, potentially disrupting the synchrony between plant phenology and breeding phenology in herbivores. A negative relationship between offspring survival in West Greenland caribou and the timing of vegetation emergence was the first finding of such a mismatch in Arctic mammals. However, other studies indicate that the energy for foetal growth and early lactation is predominantly drawn from stored energy reserves typical of ‘capital’ breeders, suggesting that conditions well before spring influence calf production more than the timing of spring onset. Here we use 20 years of observations of marked Svalbard reindeer to evaluate determinants of annual recruitment, as measured by the presence of a calf at foot in mid-summer. Spring temperatures and the enhanced vegetation index were used as proxies for spring onset, while data on body mass and pregnancy rates in late winter allowed us to determine maternal condition and the reproductive status before spring. Pregnancy rate, offspring survival and annual recruitment were all strongly correlated with average late winter adult female body mass (r = 0.87; r = 0.83; r = 0.92, respectively). Contrary to the findings in West Greenland, neither early calf survival nor annual recruitment were correlated with the two measures of annual variation in spring phenology (r = – 0.07, p = 0.8 and r = – 0.15, p = 0.6, respectively). We also revisit the Greenland data and reveal that the pattern of covariance between early and late measures of fecundity, as well as between early measures of fecundity and offspring survival, correspond with the results from Svalbard. Our results emphasize that conditions affecting maternal body mass during winter explain close to all the variation in recruitment, questioning the importance of the role of a mismatch between plant phenology and calving date.

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., Extreme climate events can cause population crashes and may threaten population persistence. Here, the authors model reindeer population dynamics and find that more frequent extremely icy winters can actually reduce extinction risk due to density dependence and a demographic shift to resilient ages.

Arctic winters have become increasingly warmer and rainier. Where permafrost prevails, winter rain (or rain-on-snow; ROS) is known to occasionally cause extensive ice layers at the snow/ground interface, i.e. "basal ice" or "ground-ice", with potentially large ecological and socio-economic implications. However, an overall lack of field data has so far restricted our predictive understanding of the environmental conditions shaping spatiotemporal variation in basal ice. Here, we use time-series of spatially replicated snowpack measurements from coastal (Ny-Ålesund area; 2000-2017) and central Spitsbergen (Nordenskiöld Land; 2010-2017), Svalbard, to analyze spatiotemporal patterns in basal ice and how they are linked with topography, weather, snowpack and climate change. As expected, both the spatial occurrence and thickness of basal ice increased strongly with the annual amount of winter rain. This effect was modified by accumulated snowfall; a deeper snowpack restricts ice formation following a minor rain event, but enhances ice formation following heavy rain due to an increased contribution of snowmelt. Accordingly, inter-annual variation in snow depth was negatively related to basal ice thickness. Annual fluctuations in basal ice thickness were strongly correlated in space (average correlation ρ = 0.40; 0-142 km distance between plots) due to strong spatial correlation in winter rain (ρ = 0.62; 14-410 km distance between meteorological stations). Models of basal ice based on meteorological time-series (1957-2017) suggested that ice-free winters (i.e. mean basal ice < 0.1 cm) had virtually not occurred since 1998, whereas such winters previously (1957-1998) occurred every three-four years on average. This detected cryosphere regime shift was linked to a parallel climate regime shift with increased winter rain amounts. Svalbard is regarded a bellwether for Arctic winter climate change. Our empirical study may therefore provide an early warning of future changes in high-arctic snowpacks. © 2019 The Author(s). Published by IOP Publishing Ltd under a CC BY 3.0 licence.

Rangifer (caribou/reindeer) management has been suggested to mitigate the temperature-driven transition of Arctic tundra into a shrubland state, yet how this happens is uncertain. Here we study this much focused ecosystem state transition in riparian areas, where palatable willows (Salix) are dominant tall shrubs and highly responsive to climate change. For the state transition to take place, small life stages must become tall and abundant. Therefore we predicted that the performance of small life stages (potential recruits) of the tall shrubs were instrumental to the focal transition, where Rangifer managed at high population density would keep the small-stage shrubs in a "browse trap" independent of summer temperature. We used a large-scale quasi-experimental study design that included real management units that spanned a wide range of Rangifer population densities and summer temperatures in order to assess the relative importance of these two driving variables. Ground surveys provided data on density and height of the small shrub life stages, while the distributional limit (shrubline) of established shrublands (the tall shrub life stage) was derived from aerial photographs. Where Rangifer densities were above a threshold of approximately 5 animals/km2 , we found, in accordance with the expectation of a "browse trap," that the small life stages of shrubs in grasslands were at low height and low abundance. At Rangifer densities below this threshold, the small life stages of shrubs were taller and more abundant indicating Rangifer were no longer in control of the grassland state. For the established shrubland state, we found that the shrubline was at a 100-m lower elevation in the management units where Rangifer had been browsing in summer as opposed to the migratory ranges with no browsing in summer. In both seasonal ranges, the shrubline increased 100 m per 1°C increment in temperature. Our study supports the proposal that Rangifer management within a sustainable range of animal densities can mitigate the much-focused transition from grassland to shrubland in a warming Arctic.

Peeters, B., V. Veiberg, A. Ø. Pedersen, A. Stien, R. J. Irvine, R. Aanes, B.-E. Sæther, O. Strand, and B. B. Hansen. 2017. Climate and density dependence cause changes in adult sex ratio in a large Arctic herbivore. Ecosphere 8(2):e01699. 10.1002/ecs2.1699 Variation 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 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 (ROS utero ). We show that females experiencing high ROS utero 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 ROS utero . Young females born after high ROS utero engage in reproductive events at lower body mass (about 2.5 kg less) than those born after low ROS utero . 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.

Parasitic nematodes are found in almost all wild vertebrate populations but few studies have investigated these host-parasite relationships in the wild. For parasites with free-living stages, the external environment has a major influence on life-history traits, and development and survival is generally low at sub-zero temperatures. For reindeer that inhabit the high Arctic archipelago of Svalbard, parasite transmission is expected to occur in the summer, due to the extreme environmental conditions and the reduced food intake by the host in winter. Here we show experimentally that, contrary to most parasitic nematodes, Marshallagia marshalli of Svalbard reindeer is transmitted during the Arctic winter. Winter transmission was demonstrated by removing parasites in the autumn, using a novel delayed-release anthelmintic bolus, and estimating re-infection rates in reindeer sampled in October, February and April. Larval stages of nematodes were identified using molecular tools, whereas adult stages were identified using microscopy. The abundance of M. marshalli adult worms and L4s increased significantly from October to April, indicating that reindeer were being infected with L3s from the pasture throughout the winter. To our knowledge, this study is the first to experimentally demonstrate over-winter transmission of a gastro-intestinal nematode parasite in a wild animal. Potential mechanisms associated with this unusual transmission strategy are discussed in light of our knowledge of the life-history traits of this parasite.

Summary 1. Input of external subsidies in the Arctic may have substantial effects on predator populations that otherwise would have been limited by low local primary productivity. 2. We explore life-history traits, age-specific fecundity, litter sizes and survival, and the population dynamics of an Arctic fox (Vulpes lagopus) population to explore the influence of the spatial distribution and temporal availability of its main prey; including both resident and migrating (external) prey resources. 3. This study reveals that highly predictable cross-boundary subsidies from the marine food web, acting through seasonal access to seabirds, sustain larger local Arctic fox populations. Arctic fox dens located close to the coast in Svalbard were found to have higher occupancy rates, as expected from both high availability and high temporal and spatial predictability of prey resources (temporally stable external subsidies). Whereas the occupancy rate of inland dens varied between years in relation to the abundance of reindeer carcasses (temporally varying resident prey). 4. With regard to demography, juvenile Arctic foxes in Svalbard have lower survival rates and a high age of first reproduction compared with other populations. We suggest this may be caused by a lack of unoccupied dens and a saturated population.

As global warming advances, there is a growing concern about the impact of extreme weather events on ecosystems. In the Arctic, more frequent unseasonal warm spells and rain-on- snow events in winter cause changes in snow-pack properties, including ground icing. Such extreme weather events are known to have severe effects across trophic levels, for instance, causing die-offs of large herbivores. However, the extent to which individuals and populations are able to buffer such events through behavioral plasticity is poorly understood. Here, we analyze responses in space use to rain-on- snow and icing events, and their fitness correlates, in wild reindeer in high-Arctic Svalbard. Range displacement among GPS-collared females occurred mainly in icy winters to areas with less ice, lower over-winter body mass loss, lower mortality rate, and higher subsequent fecundity, than the departure area. Our study provides rare empirical evidence that mammals may buffer negative effects of climate change and extreme weather events by adjusting behavior in highly stochastic environments. Under global warming, behavioral buffering may be important for the long-term population persistence in mobile species with long generation time and therefore limited ability for rapid evolutionary adaptation. Arctic; climate; GPS; ice; ideal-free distribution; migration; movement; Rangifer tarandus platyrhynchus; space use; Svalbard; Svalbard reindeer; time-to-event analysis. Attribution 3.0 Unported (CC BY 3.0)

The cumulative effects of climate warming on herbivore vital rates and population dynamics are hard to predict, given that the expected effects differ between seasons. In the Arctic, warmer summers enhance plant growth which should lead to heavier and more fertile individuals in the autumn. Conversely, warm spells in winter with rainfall (rain-on-snow) can cause ‘icing’, restricting access to forage, resulting in starvation, lower survival and fecundity. As body condition is a ‘barometer’ of energy demands relative to energy intake, we explored the causes and consequences of variation in body mass of wild female Svalbard reindeer (Rangifer tarandus platyrhynchus) from 1994 to 2015, a period of marked climate warming. Late winter (April) body mass explained 88% of the between-year variation in population growth rate, because it strongly influenced reproductive loss, and hence subsequent fecundity (92%), as well as survival (94%) and recruitment (93%). Autumn (October) body mass affected ovulation rates but did not affect fecundity. April body mass showed no long-term trend (coefficient of variation, CV = 8.8%) and was higher following warm autumn (October) weather, reflecting delays in winter onset, but most strongly, and negatively, related to ‘rain-on-snow’ events. October body mass (CV = 2.5%) increased over the study due to higher plant productivity in the increasingly warm summers. Density-dependent mass change suggested competition for resources in both winter and summer but was less pronounced in recent years, despite an increasing population size. While continued climate warming is expected to increase the carrying capacity of the high Arctic tundra, it is also likely to cause more frequent icing events. Our analyses suggest that these contrasting effects may cause larger seasonal fluctuations in body mass and vital rates. Overall our findings provide an important ‘missing’ mechanistic link in the current understanding of the population biology of a keystone species in a rapidly warming Arctic. Keywords: climate change, density dependence, extreme events, icing, nutrition, primary production, Rangifer, reindeer, Svalbard, weather Attribution 4.0 International (CC BY 4.0)

Arctic Cervids face considerable challenges in sustaining life in a harsh and highly seasonal environment, and when to forage is a key component of the survival strategy. We predict that a cervid maximizes net intake of energy to change the duration of feeding-ruminating cycles depending on season, and pays no attention to light or other activity-entraining cues. Still, in periods of bad weather it may pay energetically to reduce exposure and heat loss. We investigated environmental impact on the seasonal and daily activity pattern of a food-limited, predator-free arctic deer, the Svalbard reindeer. We found that the reindeer indeed had season-dependent feeding-rumination intervals, with no distinct peaks in activity at sunrise and sunset, as would be expected if animals maximize energy intake rates in predator-free environments. However, they temporarily reduced activity when exposed to low temperature and increased precipitation during winter, possibly to conserve energy. We provide insight into the behavioural strategy of Svalbard reindeer which enables them to cope with such an extreme environment.

Despite the intensive research effort directed at predicting the effects of climate change on plants in the Arctic, the impact of environmental change on species' distributions remains difficult to quantify. Predictive habitat distribution models provide a tool to predict the geographical distribution of a species based on the ecological gradients that determine it, and to estimate how the distribution of a species might respond to environmental change. Here, we present a model of the distribution of the dwarf shrub Dryas octopetala L. around the fjord Kongsfjorden, Svalbard. The model was built from field observations, an Advanced Space-borne Thermal Emission and Reflection Radiometer (ASTER) image, a GIs database containing environmental data at a spatial resolution of 20 m, and relied on generalized linear models (GLMs). We used a logistic GLM to predict the occurrence of the species and a Gaussian GLM to predict its abundance at the sites where it occurred. Temperature and topographical exposure and inclination of a site appeared to promote both the occurrence and the abundance of D. octopetala. The occurrence of the species was additionally negatively influenced by snow and water cover and topographical exposure towards the north, whereas the abundance of the species appeared lower on calciferous substrates. Validation of the model using independent data and the resulting distribution map showed that they successfully recover the distribution of D. octopetala in the study area (kappa = 0.46, AUC =0.81 for the logistic GLM [n - 200], r(2) = 0.29 for the Gaussian GLM [n - 36]). The results further highlight that models predicting the local distribution of plant species in an Arctic environment would greatly benefit from data on the distribution and duration of snow cover. Furthermore, such data are necessary to make quantitative estimates for the impact of changes in temperature and winter precipitation on the distribution of plants in the Arctic. (C) 2005 Elsevier Inc. All rights reserved.

All Together Now Environmental drivers, such as extreme weather events, impact population dynamics and can synchronize such dynamics across populations within a species. Given that many species depend on similar resources, such events might also be expected to synchronize dynamics across species, but the complexity of multispecies communities makes it difficult to reveal potential drivers in common. Hansen et al. (p. 313 ) took advantage of the simplicity of the year-round community on the high-arctic island of Spitsbergen to test for the presence of synchrony. Population fluctuations were synchronized across the three herbivore species (Svalbard reindeer, Svalbard rock ptarmigan, and sibling vole) and the single resident predator, the arctic fox, was in lagged synchrony. The driver of these fluctuations appears to be extreme winter rain-on-snow events that reduce the availability of winter forage due to ice cover.

1. Mesocarnivores increase in number and geographic range in human-disturbed ecosystems with cascading negative impact on biodiversity. To mitigate such impacts, it is essential to identify the proximate causes of such mesocarnivore releases. Here, we assess to what extent increased partial migration in semi-domesticated tundra reindeer induce a response in boreal and arctic mesocarnivores. 2. We used a large-scale and multiyear quasi-experimental study design with camera traps deployed on coastal tundra peninsulas in northern Norway to estimate area occupancy of the whole carnivore community. These peninsulas represent summer pastures for separate semidomestic reindeer herds that, owing to different degrees of partial migration, now display spatially and temporally variable densities of year-round resident reindeer. We estimated resident reindeer density by means of aerial surveys. 3. Area occupancy of all the recorded carnivore species increased strongly when resident reindeer densities exceeded 1 5 deer km 2. 4. Most of the increasing carnivore species were typical boreal forest species, implying range expansions into tundra when provided with stable food resources (prey and carrion) in terms of resident reindeer. 5. Synthesis and applications. We found that boreal mesocarnivores, known to negatively impact the productivity of reindeer and arctic wildlife of conservation concern, steeply increased in tundra areas with many year-round resident reindeer due to increased partial migration. To avoid such negative impacts, actions should be taken to minimize residency in tundra reindeer.

Global warming is expected to cause earlier springs and increased primary productivity in the Arctic. These changes may improve food availability for Arctic herbivores, but may also have negative effects by generating a mismatch between the surge of high quality food in the spring and the timing of reproduction. We analyzed a 10 year dataset of satellite derived measures of vegetation green-up, population densities, calf body masses and female reproductive success in 19 reindeer (Rangifer tarandus) populations in Northern Norway. An early onset of spring and high peak plant productivity had positive effects on calf autumn body masses and female reproductive success. In addition, body masses and reproductive success were both negatively related to population density. The quantity of food available, as determined by the onset of vegetation green-up and plant productivity over the summer were the main drivers of body mass growth and reproductive success. We found no evidence for an effect of the speed of spring green-up. Nor did we detect a negative mismatch between early springs and subsequent recruitment. Effects of global warming on plant productivity and onset of spring is likely to positively affect sub-Arctic reindeer. Population densities, vegetation green-up, and plant productivity: impacts on reproductive success and juvenile body mass in reindeer

Assessing the role of weather in the dynamics of wildlife populations is a pressing task in the face of rapid environmental change. Rodents and ruminants are abundant herbivore species in most Arctic ecosystems, many of which are experiencing particularly rapid climate change. Their different life-history characteristics, with the exception of their trophic position, suggest that they should show different responses to environmental variation. Here we show that the only mammalian herbivores on the Arctic islands of Svalbard, reindeer ( Rangifer tarandus ) and sibling voles ( Microtus levis ), exhibit strong synchrony in population parameters. This synchrony is due to rain-on-snow events that cause ground ice and demonstrates that climate impacts can be similarly integrated and expressed in species with highly contrasting life histories. The finding suggests that responses of wildlife populations to climate variability and change might be more consistent in Polar regions than elsewhere owing to the strength of the climate impact and the simplicity of the ecosystem.

International audience; Echinococcus multilocularis is a threatening cestode involved in the human alveolar echinococcosis. The parasite, mainly described in temperate regions of the Northern hemisphere was described for the first time in 1999 in the High Arctic Svalbard archipelago, Norway. The origin of this contamination could be due to an anthropogenic introduction from mainland Europe by domestic dogs or with the introduction of the sibling vole, perhaps from mainland Russia (St. Petersburg area), or with roaming Arctic foxes, known as the main definitive host of the parasite in Arctic regions. The genetic diversity of E. multilocularis in Svalbard was investigated here for the first time by genotyping using EmsB microsatellite and compared to other genotyped populations in the main worldwide endemic areas. We found low polymorphism amongst the 27 metacestode isolates from sibling voles trapped in the core of the distribution area of the vole on Svalbard. E. mutilocularis Arctic populations, using the Arctic fox as the definitive host, were genetically separated from European temperate populations that use the red fox, but closely related to St. Lawrence Island samples from Alaska. The result is inconsistent with the hypothesis of an anthropogenic introduction from mainland Europe, but can be seen as consistent with the hypothesis that Arctic foxes introduced E. multilocularis to Svalbard.

Despite numerous studies of how climate change may affect life history of mammals, few have documented the direct impact of climate on behavior. The Arctic is currently warming, and rain-on-snow and thaw–freeze events leading to ice formation on the ground may increase both in frequency and spatial extent. This is in turn expected to be critical for the winter survival of arctic herbivores. Svalbard reindeer (Rangifer tarandus plathyrynchus) have small home ranges and may therefore be vulnerable to local “locked pasture” events (ice layers limit access to plant forage) due to ground-ice formation. When pastures are “locked,” Svalbard reindeer are faced with the decision of staying and live off a diminishing fat store, or trying to escape beyond the unknown spatial borders of the ice. We demonstrate that Svalbard reindeer do the latter, as icing events cause an immediate increase in range displacement between 5-day observations. Population-level responses of previous icing events may therefore not accurately predict future responses if the spatial extent of icing increases. The impact of single events may be more severe if it exceeds the maximum movement distances, so that the spatial displacement strategy reported here no longer buffers climate effects.

Samples (blood or tissue fluid) from 594 arctic foxes (Alopex lagopus), 390 Svalbard reindeer (Rangifer tarandus platyrhynchus), 361 sibling voles (Microtus rossiaemeridionalis), 17 walruses (Odobenus rosmarus), 149 barnacle geese (Branta leucopsis), 58 kittiwakes (Rissa tridactyla), and 27 glaucous gulls (Larus hyperboreus) from Svalbard and nearby waters were assayed for antibodies against Toxoplasma gondii using a direct agglutination test. The proportion of seropositive animals was 43% in arctic foxes, 7% in barnacle geese, and 6% (1 of 17) in walruses. There were no seropositive Svalbard reindeer, sibling voles, glaucous gulls, or kittiwakes. The prevalence in the arctic fox was relatively high compared to previous reports from canid populations. There are no wild felids in Svalbard and domestic cats are prohibited, and the absence of antibodies against T. gondii among the herbivorous Svalbard reindeer and voles indicates that transmission of the parasite by oocysts is not likely to be an important mechanism in the Svalbard ecosystem. Our results suggest that migratory birds, such as the barnacle goose, may be the most important vectors bringing the parasite to Svalbard. In addition to transmission through infected prey and carrion, the age-seroprevalence profile in the fox population suggests that their infection levels are enhanced by vertical transmission.

Living in the high Arctic, the Svalbard reindeer (Rangifer tarandus platyrhynchus) and its trichostrongyle nematodes experience a long cold winter from October to late May/early June. Over this period, transmission would be expected to be low. However, in culled reindeer the abundance of infection increased from autumn to late winter, providing evidence for continued transmission within this period. To our knowledge this is the first time this has been demonstrated in a climate with temperatures consistently below 0 degrees C. In one winter (1996-1997), the average fraction of nematodes found as larvae in the abomasal mucosa increased from around 10% to 50% between October and March. This suggests that arrested development took place throughout the winter. We found no evidence for an efficient acquired immune response towards the nematodes. The abundance of infection did not tend to decrease with increasing host age after an earlier peak, but levelled off instead, as predicted by a simple immigration-death model. In the late winter when the nutritional plane is low, both adult reindeer and calves had high worm burdens at intensities that may affect their condition and fitness.