Congruent responses to weather variability in high arctic herbivores

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.     

Negative density-dependent emigration of males in an increasing red deer population

In species with polygynous mating systems, females are regarded as food-limited, while males are limited by access to mates. When local density increases, forage availability declines, while mate access for males may increase due to an increasingly female-biased sex ratio. Density dependence in emigration rates may consequently differ between sexes. Here, we investigate emigration using mark-recovery data from 468 young red deer Cervus elaphus marked in Snillfjord, Norway over a 20-year period when the population size has increased sixfold. We demonstrate a strong negative density-dependent emigration rate in males, while female emigration rates were lower and independent of density. Emigrating males leaving the natal range settled in areas with lower density than expected by chance. Dispersing males moved 42 per cent longer at high density in 1997 (37 km) than at low density in 1977 (26 km), possibly caused by increasing saturation of deer in areas surrounding the marking sites. Our study highlights that pattern of density dependence in dispersal rates may differ markedly between sexes in highly polygynous species. Contrasting patterns reported in small-scale studies are suggestive that spatial scale of density variation may affect the pattern of temporal density dependence in emigration rates and distances.     

Evidence for a trade-off between early growth and tooth wear in Svalbard reindeer

Ruminants depend on efficient physical degradation of forage through chewing to increase the surface area of the food particles presented to the microflora. Fossil evidence suggests that increased molar height is an adaptation for wear tolerance in dry ecosystems with sparse vegetation, but no study has shown selection pressure for hypsodonty in contemporary ruminants. We explored the relationships between particle size in rumen, tooth wear (scanned molar occlusal topography), age and body mass of female Svalbard reindeer living in an arctic desert at 78 degrees latitude on Svalbard. We predicted that (H1) if the rumen particle size is determined mainly by constraints due to tooth wear, and if tooth wear is mainly a function of age, average particle size in rumen should increase with age. From allometric relations it is known that larger individuals can survive on a lower-quality diet, we therefore predicted (H2) larger particle sizes with increases in (ln) body mass, irrespective of age and wear. Lastly, if there is a trade-off between growth and tooth wear in dry ecosystems (a selection pressure for hypsodonty), we predicted (H3) that teeth of heavier animals should be more worn than those of lighter animals of the same age. The proportion of small particles (<1.0 mm) decreased rapidly with increasing age (consistent with H1). Heavier females within an age class had more worn teeth (consistent with H3) than lighter ones. A close-to-isometric relationship between particle size and body mass suggested that heavier animals partly compensated for reduced tooth efficiency by chewing more. We provide the first evidence of a trade-off between fast early growth and wear (a somatic cost) of a senescence-related trait--the structure and height of the molar--in a wild ruminant inhabiting an arctic desert where selection pressure for increased tooth height is expected. This suggests that foraging conditions are more extreme than the environment in which the species originally evolved.     

An integrated population model for a long‐lived ungulate: more efficient data use with Bayesian methods

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.     

Dental microwear textural analyses to track feeding ecology of reindeer: a comparison of two contrasting populations in Norway

Using dental microwear textural analysis, we aim to explore variations in feeding habits within and between wild reindeer populations (Rangifer tarandus) in Norway. Specimens from the two wild reindeer areas of Knutshø and Hardangervidda were hunted between late August and the end of September 2014, and found to share very similar topological and ecological characteristics (continental alpine tundra). Despite the homogenisation of plant resources at the end of the snow-free period, dental microwear textural analysis on cheek teeth indicates differences in feeding behaviors between the two populations, in that reindeer from Knutshø might display a more “grazing” signal than the ones from Hardangervidda. Such differences in dental microwear textures reflect the differences in feeding habits that could be linked with differences in population density of reindeer. The Knutshø reindeer density is lower than at Hardangervidda, where primary food resources like grasses, sedges, and lichens are less abundant. Analyses also show the differences in gender for the population of Knutshø, but more studies are necessary for the clearest interpretation of data. Our results emphasize the need to explore local variations at a monthly time frame in order to better assess the versatility of dental microwear textures before applying this kind of analysis to either extant or extinct multi-population scales.     

Contrasting effects of summer and winter warming on body mass explain population dynamics in a food‐limited Arctic herbivore

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.     

Bigger teeth for longer life? Longevity and molar height in two roe deer populations

The role of tooth wear as a proximate cause of senescence in ruminants has recently been highlighted. There are two competing hypotheses to explain variation in tooth height and wear; the diet-quality hypothesis predicting increased wear in low-quality habitats, and the life-history hypothesis predicting molar height to be related to expected longevity. We compared tooth height and wear from roe deer of known age from two contrasting populations of roe deer (Capreolus capreolus) in France: Trois Fontaines (TF) with good habitat and shorter animal life expectancy and Chizé (CH) with poor habitat and longer animal life expectancy. There was no population difference in tooth wear, leading to rejection of the diet-quality hypothesis. However, despite their smaller body size, initial molar height for animals from CH was larger than for animals from TF. This provides the first evidence that variation in longevity between populations can lead to differences in molar height within a species.     

Partial migration in expanding red deer populations at northern latitudes – a role for density dependence?

Partial migration is common in ungulates living in highly seasonal environments. Typically, at higher latitudes, this involves movement between high elevation summer areas used during breeding and lowland areas with less snow used during winter. Snow depth is regarded the main cause of migration to low elevation, but it is less clear why deer migrate to high elevation in spring. The forage maturation hypothesis explains the upward migration due to plant phenology. We here present also an alternative and non‐exclusive hypothesis, that deer migrate uphill in summer to escape competition due to the high density in winter areas (the competition avoidance hypothesis). We also suggest that social fences may play a role at high population density. Based on a unique study of 141 GPS‐marked red deer from seven regions covering the main distribution in Norway, we found that the proportion of migrants in the populations varied from 38% to 100%. Migration was more common in areas with a diverse topography, i.e. for areas with access to high elevation. Further, we found evidence that migration was negatively density dependent, and that fall migration was delayed at high density. We suggest that a combination of avoidance of competition in high density winter ranges, social fencing during summer in addition to the forage maturation and predation risk avoidance hypotheses, is needed to explain migration patterns of northern ungulates.     

Increased effect of harsh climate in red deer with a poor set of teeth

Teeth are vital for mammal performance and especially in ungulates relying on mechanical decomposition of plant material for effective microbial digestion and energy uptake. The main focus of the role of teeth in ungulate life histories has been on tooth wear, while no one has addressed to what extent deviation from the natural set of teeth (maldentition) causes variation in individual fitness components. Based on mandibles from 41,066 individual red deer (Cervus elaphus L.) collected from 1969 to 2001, we tested whether maldentition had an effect on individual body condition and whether this effect depended on environmental harshness. Females with maldentition (0.6% of the population) were in a poorer condition than individuals without tooth anomalies and the effect increased during unfavorable climatic conditions. The effect of maldentition in males was less clear. This study indicates that a well-functioning set of teeth is essential for mammal performance, and that selection pressure against (dental) anomalies is more pronounced when climate is unfavorable. Electronic Supplementary Material Supplementary material is available for this article at     

Social rank,feeding and winter weight loss in red deer: any evidence of interference competition?

During winter at northern latitudes, large herbivores often exploit patches of concentrated, relatively high quality forage, which may lead to interference competition. The factors affecting success in contests and subsequent dominance rank, such as age and body weight, remain key issues in ungulate behavioural ecology. Maternal effects on offspring body weight are well known, but few studies have investigated if mothers social rank influence offspring rank. Moreover, no study has related dominance rank in ungulates to weight loss during winter. Outcomes of social interactions (n=7,609), feeding time and spatial position in red deer (Cervus elaphus) hinds and calves, and weight loss of calves, were registered from 1981 to 1996 at six winter-feeding sites within the county of Sør-Trøndelag in Norway. The level of aggressiveness was higher among calves than among adult hinds, and the factors determining the outcome of contests also differed. The initiator won the majority of interactions (more than 90% in both hinds and calves). Social rank was related to both age and body weight in adult hinds, and related to body weight and mother rank in calves. The relationship between feeding time and rank was non-linear. Feeding time was correlated with rank only among high ranked hinds, while there was no such relationship among low ranked hinds or calves. There was no correlation between winter weight loss and social rank in calves. Our study therefore underlines that, although frequent aggression is observed at artificial feeding sites of northern herbivores, this is not necessarily sufficient to give rise to interference competition.