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Arctic ecosystems are changing dramatically with warmer and wetter conditions resulting in complex interactions between herbivores and their forage. We investigated how Svalbard reindeer (Rangifer tarandus platyrhynchus) modify their late winter diets in response to long-term trends and interannual variation in forage availability and accessibility. By reconstructing their diets and foraging niches over a 17-year period (1995-2012) using serum δ 13 C and δ 15 N values, we found strong support for a temporal increase in the proportions of graminoids in the diets with a concurrent decline in the contributions of mosses. This dietary shift corresponds with graminoid abundance increases in the region and was associated with increases in population density, warmer summer temperatures and more frequent rain-on-snow (ROS) in winter. In addition, the variance in isotopic niche positions, breadths, and overlaps also supported a temporal shift in the foraging niche and a dietary response to extreme ROS events. Our long-term study highlights the mechanisms by which winter and summer climate changes cascade through vegetation shifts and herbivore population dynamics to alter the foraging niche of Svalbard reindeer. Although it has been anticipated that climate changes in the Svalbard region of the Arctic would be detrimental to this unique ungulate, our study suggests that environmental change is in a phase where conditions are improving for this subspecies at the northernmost edge of the Rangifer distribution., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

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)'.

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.

The costs of reproduction are important in shaping individual life histories, and hence population dynamics, but the mechanistic pathways of such costs are often unknown. Female reindeer have evolved antlers possibly due to interference competition on winter-feeding grounds. Here, we investigate if variation in antler size explains part of the cost of reproduction in late winter mass of female reindeer. We captured 440 individual Svalbard reindeer a total of 1426 times over 16 years and measured antler size and body mass in late winter, while presence of a 'calf-at-heel' was observed in summer. We found that reproductive females grew smaller antlers and weighed 4.3 kg less than non-reproductive females. Path analyses revealed that 14% of this cost of reproduction in body mass was caused by the reduced antler size. Our study is therefore consistent with the hypothesis that antlers in female Rangifer have evolved due to interference competition and provides evidence for antler growth as a cost of reproduction in females. Antler growth was constrained more by life history events than by variation in the environment, which contrasts markedly with studies on male antlers and horns, and hence increases our understanding of constraints on ornamentation and life history trade-offs.

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.

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., (Copyright © 2012 Australian Society for Parasitology Inc. Published by Elsevier Ltd. All rights reserved.)

The intestinal parasite community of Arctic foxes (Vulpes lagopus) on the Svalbard archipelago in the High Arctic was investigated in relation to the abundance and distribution of intermediate hosts. Five species of cestodes (Echinococcus multilocularis, Taenia crassiceps, Taenia polyacantha, Taenia krabbei and Diphyllobothrium sp.), ascaridoid nematodes and one unidentified acanthocephalan species were found. The cestodes E. multilocularis, T. crassiceps and T. polyacantha all showed a decreasing prevalence in the fox population with increasing distance from their spatially restricted intermediate host population of sibling voles (Microtus levis). In addition, the prevalence of E. multilocularis in a sample from the vole population was directly related to the local vole abundance. The cestode T. krabbei uses reindeer as intermediate host, and its prevalence in female foxes was positively related to the density of reindeer (Rangifer tarandus platyrhyncus). Finally, the prevalence of the ascaridoid nematodes also decreased with increasing distance from the vole population, a finding that is consistent with the idea that voles are involved in transmission, most likely as paratenic hosts. The prevalence of the remaining species (Diphyllobothrium sp. and an unidentified acanthocephalan) was very low. We conclude that the distribution and abundance of intermediate host structure the gastrointestinal parasite community of the Arctic fox on the Svalbard archipelago.

In Svalbard, Norway, the only intermediate host for Echinococcus multilocularis, the sibling vole, has restricted spatial distribution. A survey of feces from the main host, the arctic fox, showed that only the area occupied by the intermediate host is associated with increased risk for human infection.

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.