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Harvesting can magnify the destabilising effects of environmental perturbations on population dynamics and, thereby, increase extinction risk. However, population-dynamic theory predicts that impacts of harvesting depend on the type and strength of density-dependent regulation. Here, we used logistic population growth models and an empirical reindeer case study to show that low to moderate harvesting can actually buffer populations against environmental perturbations. This occurs because of density-dependent environmental stochasticity, where negative environmental impacts on vital rates are amplified at high population density due to intra-specific resource competition. Simulations from our population models show that even low levels of harvesting may prevent overabundance, thereby dampening population fluctuations and reducing the risk of population collapse and quasi-extinction following environmental perturbations. Thus, depending on the species' life history and the strength of density-dependent environmental drivers, low to moderate harvesting can improve population resistance to increased climate variability and extreme weather expected under global warming.

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.

The long-term project on St Kilda, including field assistant Jill Pilkington, has been largely funded by the UK Natural Environment Research Council and for one period by the Wellcome Trust. 1. The population of Soay sheep on the island of Hirta in the Outer Hebrides has been the subject of continuous study for more than 35 years. This paper focuses on the botanical aspects of the plant-herbivore interaction, showing how the vegetation affects and is affected by the sheep. 2. Grazing impacts on biomass and spatial structure varied across plant communities, with Holcus/Agrostis grasslands affected most and Wet Heath least, consistent with the hypothesis that herbivore impacts are proportional to plant productivity. 3. Within plant communities, the negative relationships between sheep numbers and plant abundance (sward height, gap/tussock cover and biomass) were significant in March but not significant in August, as expected if sheep numbers are limited by food supply in winter. 4. In most species, flower stem density declined with increasing sheep numbers. There were no examples where unpalatable plants showed increased flowering (e.g. from competitor release under selective grazing). 5. Plant production in temporary grazing exclosures (above ground net primary production) was greatest in Holcus/Agrostis grassland (12.6t/ha dry matter/year), lower in Nardus grassland (5.1t/ha/yr) and least in Plantago sward (1.3t/ha/yr) associated with differences in historical nutrient supply and microclimate. 6. The net effect of grazing on plant species richness was positive: a few highly palatable species were excluded, but small-scale coexistence of grazing tolerant species was enhanced by defoliation. 7. The Soay sheep population fluctuated from 908 (in 1988) to 2208 (in 2009), increasing by an average of 39 extra animals per year over the period 1985-2011. Between 2011 and 2020, the population fluctuated less widely and showed no trend. 8. Population change (ln(N(t+1)/N(t)) was inversely density dependent but positively correlated with plant production in Holcus/Agrostis grassland which increased during the study. 9. This plant-herbivore interaction is highly resilient, and though some species (Festuca rubra and Ranunculus acris) declined in the Holcus/Agrostis grassland, there was no significant upward trend in the abundance of unpalatable plant species. 10. Synthesis. Implications for future studies and analyses of plant-herbivore data. Concentrating on estimating primary productivity and herbivore offtake, rather than simply measuring change in plant biomass, is likely to provide greatly improved explanatory power for understanding herbivore population dynamics. Postprint

The ‘Moran effect’ predicts that dynamics of populations of a species are synchronized over similar distances as their environmental drivers. Strong population synchrony reduces species viability, but spatial heterogeneity in density dependence, the environment, or its ecological responses may decouple dynamics in space, preventing extinctions. How such heterogeneity buffers impacts of global change on large‐scale population dynamics is not well studied. Here, we show that spatially autocorrelated fluctuations in annual winter weather synchronize wild reindeer dynamics across high‐Arctic Svalbard, while, paradoxically, spatial variation in winter climate trends contribute to diverging local population trajectories. Warmer summers have improved the carrying capacity and apparently led to increased total reindeer abundance. However, fluctuations in population size seem mainly driven by negative effects of stochastic winter rain‐on‐snow (ROS) events causing icing, with strongest effects at high densities. Count data for 10 reindeer populations 8–324 km apart suggested that density‐dependent ROS effects contributed to synchrony in population dynamics, mainly through spatially autocorrelated mortality. By comparing one coastal and one ‘continental’ reindeer population over four decades, we show that locally contrasting abundance trends can arise from spatial differences in climate change and responses to weather. The coastal population experienced a larger increase in ROS, and a stronger density‐dependent ROS effect on population growth rates, than the continental population. In contrast, the latter experienced stronger summer warming and showed the strongest positive response to summer temperatures. Accordingly, contrasting net effects of a recent climate regime shift—with increased ROS and harsher winters, yet higher summer temperatures and improved carrying capacity—led to negative and positive abundance trends in the coastal and continental population respectively. Thus, synchronized population fluctuations by climatic drivers can be buffered by spatial heterogeneity in the same drivers, as well as in the ecological responses, averaging out climate change effects at larger spatial scales., Spatially autocorrelated weather and climate may cause population co‐fluctuations over large distances. We show that increasingly frequent rain‐on‐snow (ROS) and icing events in winter synchronize the annual dynamics of Svalbard reindeer populations, while, paradoxically, spatial variation in ROS trends and density‐dependent weather effects cause diverging local population trajectories in the long run. Such decoupling of population dynamics increases species viability under a rapidly warming high‐Arctic climate.

The fasting endurance hypothesis (FEH) predicts strong selection for large body size in mammals living in environments where food supply is interrupted over prolonged periods of time. The Arctic is a highly seasonal and food restricted environment, but contrary to predictions from the FEH, empirical evidence shows that Arctic mammals are often smaller than their temperate conspecifics. Intraspecific studies integrating physiology and behaviour of different‐sized individuals, may shed light on this paradox. We tested the FEH in free‐living Svalbard reindeer (Rangifer tarandus platyrhynchus). We measured daily energy expenditure (DEE), subcutaneous body temperature (Tsc) and activity levels during the late winter in 14 adult females with body masses ranging from 46.3 to 57.8 kg. Winter energy expenditure (WEE) and fasting endurance (FE) were modelled dynamically by combining these data with body composition measurements of culled individuals at the onset of winter (14 years, n = 140) and variation in activity level throughout winter (10 years, n = 70). Mean DEE was 6.3±0.7 MJ day−1. Lean mass, Tsc and activity had significantly positive effects on DEE. Across all 140 individuals, mean FE was 85±17 days (range 48–137 days). In contrast to the predictions of the FEH, the dominant factor affecting FE was initial fat mass, while body mass and FE were not correlated. Furthermore, lean mass and fat mass were not correlated. FE was on average 80% (45 days) longer in fat than lean individuals of the same size. Reducing activity levels by ~16% or Tsc by ~5% increased FE by 7%, and 4%, respectively. Our results fail to support the FEH. Rather, we demonstrate that (i) the size of fat reserves can be independent of lean mass and body size within a species, (ii) ecological and environmental variation influence FE via their effects on body composition, and (iii) physiological and behavioural adjustments can improve FE within individuals. Altogether, our results suggest that there is a selection in Svalbard reindeer to accumulate body fat, rather than to grow structurally large. he Arctic, activity, biologging, daily energy expenditure, doubly labelled water, intraspecific scaling, subcutaneous body temperature, Svalbard reindeer to accumulate body fat, rather than to grow structurally large.

Harvesting can magnify the destabilizing effects of environmental perturbations on population dynamics and, thereby, increase extinction risk. However, population-dynamic theory predicts that impacts of harvesting depend on the type and strength of density-dependent regulation. Here, we used population models for a range of life histories and an empirical reindeer case study to show that harvesting can actually buffer populations against environmental perturbations. This occurs because of density-dependent environmental stochasticity, where negative environmental impacts on vital rates are amplified at high population density due to intra-specific resource competition. Simulations from our population models show that even low levels of proportional harvesting may prevent overabundance, thereby dampening population fluctuations and reducing the risk of population collapse and quasi-extinction induced by environmental perturbations. Thus, depending on the species’ life history and the strength of density-dependent environmental drivers, harvesting can improve population resistance to increased climate variability and extreme weather expected under global warming.

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.

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.

Early-life environmental conditions may generate cohort differences in individual fitness, subsequently affecting population growth rates. Three, nonmutually exclusive hypotheses predict the nature of these fitness differences: (1) silver spoon effects, where individuals born in good conditions perform better across the range of adult environments; (2) the “environmental saturation” hypothesis, where fitness differences only occur in intermediate adult environmental conditions; and (3) the “environmental matching” or “predictive adaptive response” (PAR) hypothesis, where fitness is highest when adult environmental conditions match those experienced in early life. We quantified the context-dependent effect of early-life environment on subsequent reproductive success, survival, and population growth rate (k) of Svalbard reindeer, and explored how well it was explained by the three hypotheses. We found that good early-life conditions increased reproductive success compared to poor early-life conditions, but only when experiencing intermediate adult environmental conditions. This is the first example of what appears to be both “beneficial” and “detrimental environmental saturation” in a natural system. Despite weak early-life effects on survival, cohorts experiencing good early-life conditions contributed to higher population growth rates, when simulating realistic variation in adult environmental conditions. Our results show how the combination of a highly variable environment and biological constraints on fitness components can suppress silver spoon effects at both extremes of the adult environmental gradient. beneficial saturation; cohort; delayed environmental effect; detrimental saturation; environmental matching; fitness; predictive adaptive response; reindeer; silver spoon.

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.

Environmental variation can generate life-long similarities among individuals born in the same breeding event, so-called cohort effects. Studies of cohort effects have to account for the potentially confounding effects of current conditions (observation year) and age of individuals. However, estimation of such models is hampered by inherent collinearity, as age is the difference between observation year (period) and cohort year. The difficulties of separating linear trends in any of the three variables in Age-Period-Cohort (APC) models are the subject of ongoing debate in social sciences and medicine but have remained unnoticed in ecology. After reviewing the use of APC models, we investigate the consequences of model specification on the estimation of cohort effects, using both simulated data and empirical data from a long-term individual-based study of reindeer in Svalbard. We demonstrate that APC models are highly sensitive to the model's treatment of age, period and cohort, which may generate spurious temporal trends in cohort effects. Avoiding grouping ages and using environmental covariates believed to be drivers of temporal variation reduces the APC identification problem. Nonetheless, ecologists should use caution, given that the specification issues in APC models may have substantial impacts on estimated effect sizes and therefore conclusions.

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.

Studies of locomotor activity in Svalbard reindeer reported the temporary absence of diel rhythms under Arctic photic conditions. However, using Lomb-Scargle periodogram analyses with high statistical power we found diel or circadian rhythmicity throughout the entire year in measures of behaviour, temperature in the rumen and heart rate in free-living Svalbard reindeer. Significant diel rhythmicity was only lacking during some of the 15-day intervals analysed in the less frequently measured heart rate. During Polar Night these rhythms were free-running and attenuated. During continual daylight in summer, rhythms where entrained to 24 hours corresponding with the daily variation in the intensity of solar radiation, but attenuated when continuous daylight coincided with the period of growing forage. Diel rhythmicity was reduced during this short period of peak foraging activity, which coincided with peak heart rate and temperature in the rumen, most likely to facilitate fattening when food is abundant. For the rest of the year, heart rate and temperature showed the most pronounced and long-lasting suppression ever found in ungulates. The profound seasonal changes in foraging, metabolic activity, and power of diel and circadian rhythmicity of Svalbard reindeer can be viewed as adaptations to the extreme living conditions in the High Arctic.

We tested whether Svalbard reindeer (Rangifer tarandus platyrhynchus) minimise the risk of gastro- intestinal nematode infection by avoiding patches with a high density of faeces. This experiment was performed in preferred summer foraging habitat. The possibility that reindeer assess infection risk on the basis of faecal contamination levels across plant communities was determined by measuring the distribution of faeces in seven plant communities, and nematode developmental success in two plant communities with contrasting soil moisture content. We explored whether variation within individual reindeer in the levels of infection by gastro-intestinal nematodes was related to their diet. Reindeer avoided pastures where faecal contamination was increased, and thereby potentially reduced the risk of becoming infected by Trichostrongyle nematodes. Dung density was inversely related to soil moisture content, with high densities of faeces in dry plant communities and low densities in wet communities. However, nematode developmental success was positively related to soil moisture content, and was highest in the wetter sites. Thus, by avoiding dry areas with high dropping densities, reindeer would tend to feed in wetter areas where nematodes thrive. Therefore, dung density may be an unreliable predictor of the risk of infection. The absence of a strong relationship between an individual’s infection level and its diet might be due to the unpredictability of pasture infection level.

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.

Capsule: Common Guillemots Uria aalge show delayed breeding and marked age-related changes in reproductive success consistent with improved performance with experience. Aims: To determine age of first breeding and age-related effects on breeding phenology and success of Common Guillemots. Methods: Resighting data from a long-term colour-ringing study of Common Guillemot chicks were combined with observations of breeding phenology and success to follow the recruitment process, breeding phenology and success of 62 birds at a major North Sea colony over a 30-year period. Results: The median age of first breeding of Common Guillemots was 6.6 years. There were no detectable costs of first breeding on return rates or the likelihood of breeding the next season but first time breeders bred later and less successfully. Age of first breeding and lifetime breeding success both varied among individuals but there was no clear optimal age of first breeding and early first breeding was not associated with higher lifetime breeding success. Conclusions: Common Guillemots in the Isle of May population delayed breeding for 3–4 years beyond physiological maturity. The marked increase in breeding success with age was consistent with improved performance with experience rather than selection for higher quality individuals. Findings from this study will inform population models by providing improved estimates of age of first breeding and age-related changes in reproductive performance.

Conventionally, small populations living on islands are expected to lose genetic variation by drift. Fluctuations in population size, combined with polygynous mating systems, are expected to contribute to the process by increasing sampling effects on genetic variation. However, in individually monitored populations of Red deer on Rum and Soay sheep on St. Kilda, which experience fluctuations in population size, two processes have been identified which mitigate loss of genetic variation. First, in a number of examples, population reductions are associated with selection. Selection may be in favour of heterozygotes, or, as we have documented in several cases, it may fluctuate in direction temporally. Second, in Soay sheep, in which mortality over population crashes is male-biased, ostensibly leading to low effective numbers of males, molecular studies show that there are systematic changes in the reproductive success of young males, and in variance in male success, that broaden genetic representation compared with expectation.

The fitness consequences of inbreeding and outbreeding are poorly understood in natural populations. We explore two microsatellite-based variables, individual heterozygosity (likely to correlate with recent inbreeding) and a new individual-specific internal distance measure, mean d2 (focusing on events deeper in the pedigree), in relation to two measures of fitness expressed early in life, birth weight and neonatal survival, in 670 red deer calves (Cervus elaphus) born on the Isle of Rum between 1982 and 1996. For comparison, we also analyse inbreeding coefficients derived from pedigrees in which paternity was inferred by molecular methods. Only 14 out of 231 calves (6.1%) had non-zero inbreeding coefficients, and neither inbreeding coefficient nor individual heterozygosity was consistently related to birth weight or neonatal survival. However, mean d2 was consistently related to both fitness measures. Low mean d2 was associated with low birth weight, especially following cold Aprils, in which foetal growth is reduced. Low mean d2 was also associated with low neonatal survival, but this effect was probably mediated by birth weight because fitting birth weight to the neonatal survival model displaced mean d2 as an explanatory variable. We conclude that in the deer population fitness measures expressed early in life do not show evidence of inbreeding depression, but they do show evidence of heterosis, possibly as a result of population mixing. We also demonstrate the practical problems of estimating inbreeding via pedigrees compared with a direct marker-based estimate of individual heterozygosity. We suggest that, together, individual heterozygosity and mean d2, estimated using microsatellites, are useful tools for exploring inbreeding and outbreeding in natural population.

Life-history theory predicts that the vital rates that influence population growth the most should be buffered against environmental fluctuations due to selection for reduced variation. However, it remains unclear whether populations actually are influenced by such “demographic buffering,” because variation in vital rates can be compared on different measurement scales, and there has been little attempt to investigate whether the choice of scale influences the chance of detecting demographic buffering. We compared two statistical approaches to examine whether demographic buffering has influenced vital rates in wild Svalbard reindeer. To account for statistical variance constraints on vital rates limited between 0 and 1 in analyses of demographic buffering, one approach is to scale observed variation by the maximum possible variation on the arithmetic scale. When applying this approach, the results suggested that demographic buffering was occurring. However, when we applied an alternative approach that identified statistical variance constraints on the logit scale, there was no evidence for demographic buffering. Thus, the choice of measurement scale must be carefully considered before one can fully understand whether demographic buffering influences life histories. Defining the appropriate scale may require an understanding of the mechanisms through which demographic buffering may have evolved. © 2016 by the Ecological Society of America. This is the authors’ accepted and refereed manuscript to the article.

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)

Environmental conditions experienced during early growth and development markedly shape phenotypic traits. Consequently, individuals of the same cohort may show similar life-history tactics throughout life. Conditions experienced later in life, however, could fine-tune these initial differences, either increasing (cumulative effect) or decreasing (compensatory effect) the magnitude of cohort variation with increasing age. Our novel comparative analysis that quantifies cohort variation in individual body size trajectories shows that initial cohort variation dissipates throughout life, and that lifetime patterns change both across species with different paces of life and between sexes. We used longitudinal data on body size (mostly assessed using mass) from 11 populations of large herbivores spread along the “slow-fast” continuum of life histories. We first quantified cohort variation using mixture models to identify clusters of cohorts with similar initial size. We identified clear cohort clusters in all species except the one with the slowest pace of life, revealing that variation in early size is structured among cohorts and highlighting typological differences among cohorts. Growth trajectories differed among cohort clusters, highlighting how early size is a fundamental determinant of lifetime growth patterns. In all species, among-cohort variation in size peaked at the start of life, then quickly decreased with age and stabilized around mid-life. Cohort variation was lower in species with a slower than a faster pace of life, and vanished at prime age in species with the slowest pace of life. After accounting for viability selection, compensatory/catch-up growth in early life explained much of the decrease in cohort variation. Females showed less phenotypic variability and stronger compensatory/catch-up growth than males early in life, whereas males showed more progressive changes throughout life. These results confirm that stronger selective pressures for rapid growth make males more vulnerable to poor environmental conditions early in life and less able to recover after a poor start. Our comparative analysis illustrates how variability in growth changes over time in closely related species that span a wide range on the “slow-fast” continuum, the main axis of variation in life-history strategies of vertebrates. This article is protected by copyright. All rights reserved.

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)

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. Grazing has been implicated in the decline of heather-dominated moorlands in Britain, but there has been little consideration of the effects of different species of herbivores on plant diversity. Sheep stocks have recently decreased in Scotland, and quantification of how this affects biodiversity is essential for understanding how different grazing regimes modify upland habitats. 2. We investigated the effects of grazing on plant biodiversity in heather/grass mosaics at 16 upland sites in Scotland. Red deer Cervus elaphus L. were present at all sites, but sheep Ovis aries L. had been removed from half of the sites. Our experimental design incorporated replication at three spatial scales, from a landscape level down to 10 × 10 m plots. 3. We quantified the relative effects of different herbivore species, vegetation structure and rainfall on heather utilisation, species richness and evenness (alpha diversity; Shannon–Wiener index) and beta diversity. 4. At all spatial scales, deer dung counts were higher and heather was shorter when sheep were absent. Furthermore, utilisation of heather was positively correlated with the amount of deer dung, the amount of grass present and smooth grass height. 5. Alpha diversity was consistently positively correlated with the relative amount of grass, but was also positively related to the amount of sheep dung at the largest spatial scale. At the finest scale, alpha diversity was negatively correlated with the amount of deer dung. Beta diversity was higher when sheep were present at all scales. 6. Synthesis and applications. Mixed grazing by sheep and deer appears to be beneficial for increasing both alpha and beta diversity and minimizing damage to heather in the uplands. The absence of sheep is likely to result in expanding deer populations and greater impact on heather. Management of grazing herbivores is an important tool for maintaining biodiversity in many ecosystems. Our results indicate that reducing livestock may alter the impacts of wild grazers on their habitats and drive changes in diversity, whereas mixed grazing can enhance habitat quality and maintain plant diversity. Therefore, this effect should be considered when devising policy recommendations.

Summary 1. In species inhabiting a seasonal environment, timing of birth may be highly correlated with fitness. However, we know little about the relative effects of conception date and gestation length in determining birth date in wild mammalian populations. 2. Here, we tested for variation in gestation length using long-term data from a wild ungulate population (red deer, with observations running from 1971 to 2007). High March temperatures during gestation were associated with shorter average gestation length (a decrease of 0·77 days for every degree warmer), but we found no significant correlations with female age, recent reproductive history or offspring sex. 3. We also tested whether early or late conceiving females adjusted gestation length to give birth closer to an optimum time. Simply fitting conception date as a fixed effect in a model of gestation length is not statistically sound as errors in determining conception date could drive an observed negative covariance; consequently we also approached this problem using a novel technique of bivariate models. We found that late conception was indeed correlated with shorter gestation length, but this was not repeatable within females, suggesting either that the observed association may have been a statistical artefact or that gestation length adjustment may only be apparent on the rare occasions that females conceive outside an optimum time window. 4. Although there were no consistent differences between females in gestation length, there were differences between males in the gestation lengths of females with whom they mated. This possibly reflects a local environment effect, grouping females who were primarily in the same area during conception and gestation. 5. We conclude that observed variation in birth date is predominantly due to systematic variation in conception date as opposed to gestation length. Additionally, we highlight the broad range of reproductive tactics utilized by a number of closely related species, and consequently the difficulties inherent in cross-species comparisons of gestation length.

Although many ecological studies of wild animal populations rely on capturing individuals at least once, few have investigated either the immediate stress responses of capture, restraint, and handling, or the longer term consequences on reproductive success. In this paper, we assume that the degree of stress increase with the time taken to capture and process an animal prior to release. The influence of time since onset of capture on body temperature and blood chemistry indicators of stress was investigated in 137 adult female Svalbard reindeer ( Rangifer tarandus platyrhynchus Vrolik, 1829) and 63 calves of both sexes captured by net in late-winter 2004 and 2005. In addition, we looked for evidence of negative effects of capture stress on reproductive success in marked animals over the period from 1995 to 2006. Body temperature increased with both chase time and the time the animals were restrained. In the latter case, body temperature increased twice as fast in nonpregnant animals (0.13 °C/min) compared with pregnant animals. Cortisol, glutamate dehydrogenase, and γ-glutamyltransferase all increased with time from onset of capture. However, the relationship for glucose with time since onset of capture increased initially before declining again. We found no evidence of a lower calving success in animals captured the previous winter when compared with animals that had not been captured the previous winter. Likewise, there was no relationship between the strength of the acute stress response (measured as cortisol concentrations) and the probability of pregnant females having a calf at foot the subsequent summer.

SUMMARYMacroparasites potentially play a significant but often ignored role in the ecology and dynamics of wild ruminant populations. In the Arctic, parasites may impact on host populations by exacerbating the effects of seasonal and limited forage availability on the condition, fecundity and survival of individuals. We studied the effects of abomasal nematode parasites and warble flies, Hypoderma tarandi, on condition and pregnancy of caribou Rangifer tarandus in the Dolphin-Union herd, Nunavut, Canada. By the end of winter, female caribou over 2 years old showed a significant decrease in body weight with increasing nematode burden, and a decrease in back fat depth with increasing warble abundance. These effects were exaggerated in the non-pregnant fraction of the population. High warble larvae burdens were also associated with significantly reduced probability of being pregnant. Our research demonstrates a negative relationship between parasites and caribou condition that may have consequences for their fitness. Additionally, we discuss the possibility that muskox Ovibos moschatus share some parasite species with the caribou and could lead to elevated burdens in the sympatric host. Parasites may have been a contributory factor in a previous winter range-shift of the caribou herd and this may reflect a form of apparent competition between the two ungulate species.

Summary 1. We use structured demographic accounting to decompose the contribution to the variance in relative population growth between 1971 and 1997 in female red deer on Rum, Scotland, in terms of different fitness components and age categories. 2. During the first 10 years (1971–80) the variance in relative population growth was small (0·00246) as the population grew from 58 to 148 having been culled previously. After this the relative variance in population growth increased to 0·00575 between 1981 and 1989, and increased again to 0·00848 between 1990 and 1997. 3. In the first phase of population growth (1971–80) changes in the birth rate contributed most to the variance in relative population growth (30%), then adult winter survival (23%), with calf summer and calf winter survival contributing less than 10%. Birth rate tended to covary strongly with adult winter survival, and to a lesser extent with calf winter survival. 4. The contribution of birth rate to the variance in relative population growth diminished to less than 5% in both the periods 1981–89 and 1990–97. In contrast, the contribution of adult winter survival increased to 36% in 1980s and 40% in 1990s. At the same time, the covariation of adult winter survival with calf winter survival increased successively to reach 40% in 1990–97. 5. Changes in population age structure were small and contributed little to the variance in relative population growth (≈ 1% over the entire study). In the 1970s the 5–7-year-old group contributed most (23%) to the variance in relative population growth, though this covaried with 8–12-year-olds. In the 1980s the 8–12-year-olds contributed most (27%) and now the covariation was with older animals (> 12-year-olds). This pattern persisted in the 1990s, though there was covariation with 5–7-year-olds too. 6. Birth rate, calf winter survival and adult winter survival were all density-dependent. The strength of density dependence within each vital rate did not differ between the 1970s, 1980s and 1990s; however, there were differences in the means of these vital rates relative to population size between the three periods. 7. The implications of temporal changes in the most important vital rates, the ‘key factors’, contributing to the variance in relative population growth rate are discussed.

1. There is increasing evidence that density-independent weather effects influence life-history traits and hence the dynamics of populations of animals. Here, we present a novel statistical approach to estimate when such influences are strongest. The method is demonstrated by analyses investigating the timing of the influence of weather on the birth weight of sheep and deer. 2. The statistical technique allowed for the pattern of temporal correlation in the weather data enabling the effects of weather in many fine-scale time intervals to be investigated simultaneously. Thus, while previous studies have typically considered weather averaged across a single broad time interval during pregnancy, our approach enabled examination simultaneously of the relationships with weekly and fortnightly averages throughout the whole of pregnancy. 3. We detected a positive effect of temperature on the birth weight of deer, which is strongest in late pregnancy (mid-March to mid-April), and a negative effect of rainfall on the birthweight of sheep, which is strongest during mid-pregnancy (late January to early February). The possible mechanisms underlying these weather-birth weight relationships are discussed. 4. This study enhances our insight into the pattern of the timing of influence of weather on early development. The method is of much more general application and could provide valuable insights in other areas of ecology in which sequences of intercorrelated explanatory variables have been collected in space or in time.

Summary 1 Rangelands, produced by grazing herbivores, are important for a variety of agricultural, hunting, recreation and conservation objectives world-wide. Typically, there is little quantitative evidence regarding the magnitude of the grazing impact of different herbivores on rangeland habitats to inform their management. 2 We quantified the grazing and trampling impact of sheep, cattle, red deer Cervus elaphus, rabbits Oryctolagus cuniculus, mountain hares Lepus timidus and red grouse Lagopus lagopus on open-hill habitats in 11 areas of upland Scotland. The degradation of heather in upland Scotland Calluna vulgaris-dominated habitats, of conservation significance at a European scale, has been attributed, anecdotally, to increasing sheep and red deer populations. 3 Field indicators of habitat condition were used to generate a five-point scale of impact in vegetation polygons of seven habitats. The presence of each herbivore species was attributed on the basis of ‘signs’ of occupancy. A Bayesian regression model was used to analyse the association of herbivore species with grazing impact on plant communities, controlling for environmental attributes. 4 Overall the presence of sheep was associated with the largest increase (7/11 areas) in grazing and trampling impact of all herbivores. Cattle had the second largest impact but generally this was restricted to fewer areas and habitats than sheep. In contrast, impacts associated with wild herbivores tended to be small and only significant locally. 5 Although red deer presence was associated with a significantly lower impact than sheep, this impact increased with increasing deer density at both land-ownership and regional scales. For sheep there was little or no evidence of density dependence. 6 Synthesis and applications. The higher impact associated with sheep presence probably reflects their greater aggregation because of their limited ranging behaviour, exacerbated by sheep being herded in places convenient for land managers. Consequently, future reductions in sheep numbers as a result of reform of European Union farming policies may limit the extent of their impact, but not necessarily the local magnitude. However, reductions in sheep stocks may lead to increases in deer densities, with greater impact, particularly in heather-dominated habitats. Where habitat conservation is a priority this may well require a reduction in deer numbers.

We develop an integrated population model for Svalbard reindeer Rangifer tarandus platyrhynchus, and demonstrate how this type of model can be used to extract more information from the data and separate different sources of variability in population estimates. Our model combines individual mark–recapture data with population counts and harvesting data within a Bayesian model framework, and accounts for observation error, environmental and demographic stochasticity,and age structure. From this model we obtain annual estimates of age-specific population size, survival and fecundity. The model provides estimates of age structure at a finer scale than that found in the census data, and enables us to estimate survival for the period before calves are first caught and marked, i.e. before they enter the individual mark–recapture data. The modeling framework provides an improved approach to studying age-structured populations that are imperfectly censused and where the demography of only a sample of individuals is known. We use data from independent censuses of the same population to evaluate population estimates obtained from the model, and show that it is successful at correcting for different types of observation error. Based on our model results, we suggest that allocating resources to the collection of supplementary mark–recapture data could improve the reliability of population projections more than making regular population censuses as exhaustive as possible. Our work demonstrates how integrated Bayesian population modeling can be used to increase the amount of information extracted from collections of data, identifying and disentangling sources of variation in individual performance and population size. This represents an important step towards increasing the predictive ability of population growth models for long-lived species experiencing changes in environmental conditions and harvesting regimes. This is the pre-peer reviewed version of the following article: An integrated population model for a long-lived ungulate: more efficient data use with Bayesian methods, which has been published in final form at http://onlinelibrary.wiley.com/doi/10.1111/oik.01924/epdf. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Self-Archiving

Regulation of ungulate populations by parasites relies on establishing a density-dependent relationship between infection and vital demographic rates which may act through the effect of parasites on body condition. We examine evidence for parasite impacts in 285 red deer (Cervus elaphus) harvested during 1991 and 1992 on the Isle of Rum. In the abomasa, prevalence of nematodes was 100% and the most abundant genus observed wereOstertagiaspecies, however, mean intensity of infection was low (less than 1000) relative to other studies. Additional species, also present in low numbers, includedNematodirusspp.,Capillariaspp.,Cooperiaspp., Monieza expanza,Oesophagostomum venulosumandTrichuris ovis. Lungworm (Dictyocaulusspp.) and tissue worm (Elaphostronygylus cervi) larvae were also observed in faecal samples. There was no evidence for acquired immunity to abomasal nematodes. Despite low levels of infection, both adult male and female deer showed significant negative correlation between indices of condition (kidney fat index, dressed carcass weight and larder weight) and intensity ofOstertagiaspp. infection. However, there was no evidence that pregnancy rate in females was related to intensity of infection. For calves, there was no relationship between body condition and intensity of infection. The apparent subclinical effects of low-level parasite infection on red deer performance could alternatively be due to animals in poorer nutritional state being more susceptible to infection. Either way the results suggest that further studies of wild populations are justified, in particular where high local host densities exist or alternative ungulate hosts are present, and, where experimental treatments are tractable.

Summary 1 Sexual segregation occurs in most species of sexually dimorphic ungulates. At least five not mutually exclusive hypotheses have been formulated to explain patterns of social, habitat and spatial segregation; the indirect competition hypothesis (H1), the nutritional needs hypothesis (H2), the activity budget hypothesis (H3), the weather sensitivity hypothesis (H4), and the predation risk hypothesis (H5). 2 Each hypothesis has a unique set of predictions with respect to the occurrence of segregation, and how seasonality, density dependence and reproductive status affect sexual segregation. 3 We tested this set of predictions in order to separate the hypotheses H1–H5 for patterns of sexual segregation of the Svalbard reindeer based on 9 years data on seasonal estimates of spatial, habitat and social (i.e. grouping with their own sex) segregation in combination with resource selection functions. 4 Our results do not support that one single mechanism causes segregation. The activity budget hypothesis, the nutritional needs hypothesis and the weather sensitivity hypothesis were all partially supported, while the predation risk hypothesis was discarded for Svalbard reindeer because predators have been absent for at least 5000 years. Several mechanisms are thus interacting to explain the full-year pattern of sexual segregation and the combination of mechanisms varies among species and populations.

Resistance to parasites is believed to have a widespread influence on demographic and adaptive processes. In systems where parasites impose a fitness cost on their host, heterozygotes may be selected because they are more resistant to parasites than homozygotes. Our objective was to assess the relationships between genomewide individual heterozygosity and abomasal nematode burdens in female Svalbard reindeer (Rangifer tarandus platyrhynchus) after the effects of host age, locality, season, and year had been accounted for. Samples were obtained from 306 female reindeer that were culled and genotyped at nine microsatellite loci. Reindeer in our study populations are mainly parasitized by the gastrointestinal nematodes Ostertagia gruehneri and Marshallagia marshalli. The infection intensity of each parasite differed between subpopulations, and among host age classes, seasons and years. We found no significant relationships between abomasal worm burdens, or lumen and mucosa larvae, of either O. gruehneri or M. marshalli and individual heterozygosity (or mean d(2)) alone or in interactions with host age, locality, and year. Although we analysed one of the largest data set available to date on gastrointestinal nematodes of a wild ruminant, we used a typical data set of nine genetic neutral markers that may have had low power to detect heterozygosity-fitness correlations. We conclude that the proportion of the variance in parasite resistance explained by individual heterozygosity for neutral genetic markers is low in Svalbard reindeer and in vertebrates in general, and we suggest that the candidate-gene approach might be more fruitful for further research on gene-fitness correlations.

Summary 1. Phenotypic plasticity and microevolution represent the two processes by which phenotypic traits in a population can track environmental change. While there is a growing literature documenting microevolution in reproductive traits in naturally occurring animal populations, few studies to date have examined either between-individual variation in levels of plasticity or how selection acts on plasticity. 2. We present here mixed-effect linear models analysing changes in calving date in relation to autumn rainfall observed over a 30-year study of 2147 red deer on the Isle of Rum, Scotland. The study period is characterized by a phase of low and rising population density (up to and including 1980), followed by a phase of high and fluctuating population density (1981 to present). 3. Variation within individual females explained a population-level trend of delayed calving dates following years of high autumn rainfall. There was significant variation between females both in their average calving dates and in their individual plastic responses of calving date to autumn rainfall. 4. Females born in the low population density phase were, on average, phenotypically plastic for the calving date‐autumn rainfall relationship, and showed significant variation in plasticity. Selection favoured individuals with early average calving dates among these females. 5. Among females born at high population density, there was on average no significant plasticity for calving date, but variation in plastic responses was still present. Selection favoured females with increasingly positive plastic responses of calving date to autumn rainfall. 6. We argue that early experience of high population density affects the physiological condition of females, making an environmental response (calving early following dry autumns) in later life physiologically untenable for all but a few high quality individuals. These same few individuals also tend to be fitter and have higher reproductive success.

Estimates of the intensity and abundance of species provide essential data for ecological, evolutionary and epidemiological studies of gastrointestinal nematode communities. These estimates are typically derived from the species composition of adult males when only males have readily scorable species-specific morphological traits. Such estimation assumes that all species in the community have the same adult sex ratio. We evaluated this assumption for the trichostrongyle nematodes Ostertagia gruehneri and Marshallagia marshalli in infracommunities in Svalbard reindeer by identifying to species adult females using a polymerase chain reaction assay. The proportion of males was found to be slightly higher in O. gruehneri than in M. marshalli. Evidence for seasonal variation and density dependence in the adult sex ratio was only found for O. gruehneri. Possible demographic mechanisms for such sex ratio variation are discussed, and stochastic models that generate density-dependent sex ratios proposed. Sex ratio variation caused substantial bias in some male-based estimates of intensity of infection, while substantial and consistent bias in estimates of abundances was only evident in late winter samples. Our results suggest that estimating sex ratios can be particularly important in individual host level studies of nematode species of low abundance.

Guanacos (Lama guanicoe) are the largest native Artiodactyl in South America and the most widely distributed. In arid Patagonia, densities are low and negatively related to domestic sheep numbers in space and time consistent with interspecific competition theory. Although guanacos and domestic sheep have been described as intermediate feeders sharing food resources, no studies have been conducted to compare their diets in sympatric conditions and explore whether the potential exists for direct interspecific competition. We assessed the diet of both species across 9 different sites and 2 seasons by microhistological analysis of fecal samples. We found that (1) guanacos and sheep are generalist herbivores feeding on a wide range of plant species; (2) both are intermediate feeders able to include both monocotyledoneous and dicotyledoneous plants in their diet; (3) both are able to change their diets seasonally; and (4) food niche overlap is high, particularly in summer when food resources are more scarce than in spring. We conclude that the potential for competition between guanacos and sheep is high and could have played a major role in the demise of guanacos. Consequently, current management practices focused on maximizing sheep numbers are not compatible with the recovery of guanaco populations.

Body condition is an important determinant of ecological fitness but is difficult to measure in field studies of live animals. Live mass and subcutaneous fat are often used as proxies for body condition and related to fitness. We investigated the relationship between blood-chemistry parameters and live mass and back-fat thickness and assessed their usefulness as predictors of ecological fitness in a wild arctic ungulate population, Svalbard reindeer (Rangifer tarandus platyrhynchus). Female reindeer were sampled in late winter between 1995 and 2002 and concentrations of blood parameters were related to subsequent survival and successful calving. There was marked annual variation in all blood parameters, live mass, and back-fat thickness, reflecting variation in weather and food availability. At the individual level, variation in blood-parameter concentrations was not closely related to variation in live mass or back-fat thickness, instead reflecting shorter term nutritional status. Blood parameters could therefore provide useful additional information, enhancing the predictive power of fitness models based on live mass. The urea:creatinine ratio significantly improved adult survival models, while β-hydroxybutyric acid and creatinine concentrations were significant predictors of calving success. The applications for blood parameters in ecological investigations look promising and should be tested more widely in other field studies.

The high degree of habitat heterogeneity and habitat fragmentation in arctic ecosystems may support a high genotypic and ecotypic variability. This may buffer the survival of plant species threatened by global climate change, which affects the Arctic more strongly than other ecosystems. Here, we assessed if two morphotypes of Polygonum viviparum (characterised by different colours of their bulbils) differ in their abundance along a snowmelt gradient, if their biomass allocation patterns are influenced differentially by environmental variables, and if the temperature dependency of bulbil germination differs between morphotypes. We found slight differences in the effect of timing of snowmelt on abundance of the morphotypes, which seem to have little ecological relevance. Total biomass and biomass allocation were similar between morphotypes and were negatively correlated with soil-water content. Bulbil germination (the onset of growth of the bulbil) was assessed over a temperature range from 2 to 25°C and results indicate an earlier (maximum of 5 days) "germination" of one morphotype, but final bulbil germination (>80%) and bulbil-germination rate were similar for both types. Bulbil germination was weakly temperature dependent, with faster emergence at higher temperatures. Overall, our results could provide no convincing evidence for differences between the two examined morphotypes that could be of ecological relevance with respect to anticipated climate change in the Arctic.

Summary 1 It is well known that gastrointestinal nematodes can have a significant impact on the growth of farmed ruminants. The clinical signs of these parasites are often subtle, with production losses mainly due to reductions in host appetite and gut function. However, little is known about the impact of this widespread group of parasites on wild ruminants. We use experiments and cross-sectional data to investigate the effects of gastrointestinal nematodes on a wild host, the Svalbard reindeer. 2 Individually marked reindeer were treated for parasites. Their body mass, back fat depth and fecundity were compared with the control group one year later. The effect of treatment on worm burdens was investigated in a subsample of animals that were culled 3 and 6 months after treatment. Also, the relationship between the intensity of infection and body mass, back fat depth and host pregnancy rates was investigated in cross-sectional data from culled reindeer. 3 The anthelmintic treatment caused an increase in the body mass, back fat depth and fecundity of the reindeer. Treatment depressed the abundance of adult parasites of Ostertagia gruehneri for at least 6 months, but had no significant effect on the abundance of adults of the other dominant parasite species, Marshallagi marshalli, or the abundance of larval stages of either species. 4 In the cross-sectional study, the probability of a reindeer being pregnant in late winter was negatively related to the abundance of adult O. gruehneri when controlling for host body mass. However, no clear evidence were found for an effect of parasitism on host condition in the cross-sectional study. 5 Our experimental results show for the first time in a natural ruminant host population that gastrointestinal nematodes can have a significant effect on host condition and fecundity. However, effects of infection on host condition was not detectable in the cross-sectional study. Also, we show that the experimental effects on host condition and fecundity is most likely to be due to a negative effect of O. gruehneri, while the experimental design did not allow detection of potential negative effects of M. marshalli.

Guanacos (Lama guanicoe) are the only wild ungulate species widely distributed across the Patagonian steppe and have undergone a precipitous population decline since the introduction of domestic sheep (Ovis aries) 100 years ago. There has been speculation that sheep ranching may have played a major role in guanaco population decline after monopolising the most productive land because of competition for forage plants. Our aim in this work was to estimate guanaco abundance and account for its variation across nine different sites, two seasons and two years. We conducted over 100 ground surveys of both guanacos and sheep and estimated the availability of the preferred plant species both animal species might select in their diet. We found that (1) sheep densities were up to 23 times higher than guanaco densities in sympatric conditions, (2) at a protected site without sheep, guanaco density was one order of magnitude higher than at the rest of the sites, (3) across nine different sites, sheep densities alone accounted for around 60% of the variation in guanaco abundance, (4) guanaco densities were negatively related to both total plant cover and availability of the preferred plant species in their diet, which were both positively associated with sheep density, and (5) within-site changes in guanaco densities between seasons and years were negatively related to changes in sheep densities. Our results are consistent with predictions on interspecific competition for food resources, although we cannot rule out possible effects of other human-related activities influencing guanaco abundance. We conclude that sheep compete with guanacos for forage in arid Patagonia.