Global climate change and reindeer: effects of winter weather on the autumn weight and growth of calves

Reindeer/caribou (Rangifer tarandus), which constitute a biological resource of vital importance for the physical and cultural survival of Arctic residents, and inhabit extremely seasonal environments, have received little attention in the global change debate. We investigated how body weight and growth rate of reindeer calves were affected by large-scale climatic variability [measured by the North Atlantic Oscillation (NAO) winter index] and density in one population in central Norway. Body weights of calves in summer and early winter, as well as their growth rate (summer to early winter), were significantly influenced by density and the NAO index when cohorts were in utero. Males were heavier and had higher absolute growth than females, but there was no evidence that preweaning condition of male and female calves were influenced differently by the NAO winter index. Increasing NAO index had a negative effect on calves' body weight and growth rate. Increasing density significantly reduced body weight and growth rate of calves, and accentuated the effect of the NAO winter index. Winters with a higher NAO index are thus severe for reindeer calves in this area and their effects are associated with nutritional stress experienced by the dams during pregnancy or immediately after calving. Moreover, increased density may enhance intra-specific competition and limits food available at the individual level within cohorts. We conclude that if the current pattern of global warming continues, with greater change occurring in northern latitudes and during winter as is predicted, reduced body weight of reindeer calves may be a consequence in areas where winters with a high NAO index are severe. This will likely have an effect on the livelihood of many northern indigenous peoples, both economically and culturally.     

The relative role of winter and spring conditions: linking climate and landscape-scale plant phenology to alpine reindeer body mass

The relative importance of winter harshness and early summer foraging conditions are of prime interest when assessing the effect of global warming on Arctic and mountainous ecosystems. We explored how climate and vegetation onset (satellite-derived normalized difference vegetation index data) determined individual performance in three reindeer populations (data on 27814 calves sampled over 11 years). Snow conditions, spring temperatures and topography were the main determinants of the onset of the vegetation. An earlier onset positively affected the body mass of calves born the following autumn, while there was no significant direct negative impact of the previous winter. This study underlines the major impact of winter and spring climatic conditions, determining the spring and summer food availability, and the subsequent growth of calves among alpine herbivores.     

Density dependence in a fluctuating wild reindeer herd; maternal vs. offspring effects

Summary The Hardangervidda wild reindeer herd in Norway is the largest in Western Europe. It has fluctuated between 7000 and 32000 animals during the last 35 years. Four density-dependent effects were found: 1. A food limitation effect due to a shift in diet after overgrazing lichen on the winter range. This led to increased tooth wear and lowered body size and fat reserves. 2. A significant correlation between population density and juvenile winter survival rate. No effect on adult female survival rate was found. 3. A cohort effect. After population increase and overgrazing, recruitment was reduced by 30% and remained so after population reduction. Birth weights had increased by 30% 5 years after population reduction and the mean calving time was earlier. As a result, after population reduction weights of newborns were 40% greater at a comparable date. Neonatal survival rate was related to maternal condition during the last part of gestation which coincides with the peak winter snow accumulation. The slow increase in adult dressed body weights (DBW) after population reduction is due to the combined effects of increased tooth wear when winter range was limiting and to the cohort-generation time, so that an improvement in neonatal survival and size was first expressed in subsequent offspring cohorts. 4. An inter-generation effect. During 30 years of resource limitation, DBW decreased by 23%, birth rate was unchanged after the first peak, while fecundity increased by 15%, suggesting increased reproductive effort per unit body weight. Natural selection for increased reproductive effort by smaller females when food was limiting was suggested. Some size-effect due to hunters selecting the largest adult phenotypes was possible but not the main cause. These results do not support some earlier hypotheses about the effects of population density on size at maturity in ungulates.     

Sexual dimorphism and intercorhort variation in reindeer calf antler length is associated with density and weather

We analysed intercohort variability of live weight and antler length of 5,123 reindeer calves. We further assessed the influence of climate and density on the interannual variation in antler length, and discussed sex-specific resource allocation and response to climate variability. Antler length varied significantly among years and between sexes, with interaction between year and sex. Body weight and antler length were highly positively correlated, showed similar intercohort variability, and had a strong allometrical link, suggesting that antler length could be an equally reliable measure of calf condition as live weight. We found a relative measure of antler length (i.e. antler length corrected for the allometric effect of body mass) to be positively influenced by increasing density and May–June precipitation, and also decreasing May–June temperature. We attributed the effect of early summer weather to its influence on forage availability and quality as well as the level of parasitic insect harassment. Gender difference in both the allometric exponents and the interannual variability suggest that young males and females may have different tactics for relative resource allocation towards growth of antlers as compared to body mass. Because antlers are costly to produce, they may be an honest signal of individual quality for both sexes. However, we found gender-specific allometry, as female calves more than males appear to prioritize their antler growth over body mass, especially when resources are limited. Thus, our results suggest that environmental variation may influence the extent of sexual dimorphism in antler length. Electronic Supplementary Material Supplementary material is available for this article at     

Demographic effects of extreme winter weather in the barn owl

Extreme weather events can lead to immediate catastrophic mortality. Due to their rare occurrence, however, the long-term impacts of such events for ecological processes are unclear. We examined the effect of extreme winters on barn owl (Tyto alba) survival and reproduction in Switzerland over a 68-year period (∼20 generations). This long-term data set allowed us to compare events that occurred only once in several decades to more frequent events. Winter harshness explained 17 and 49% of the variance in juvenile and adult survival, respectively, and the two harshest winters were associated with major population crashes caused by simultaneous low juvenile and adult survival. These two winters increased the correlation between juvenile and adult survival from 0.63 to 0.69. Overall, survival decreased non-linearly with increasing winter harshness in adults, and linearly in juveniles. In contrast, brood size was not related to the harshness of the preceding winter. Our results thus reveal complex interactions between climate and demography. The relationship between weather and survival observed during regular years is likely to underestimate the importance of climate variation for population dynamics.     

Disease transmission in an extreme environment: nematode parasites infect reindeer during the Arctic winter

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.     

Age class,density and temporal effects on diet composition of sheep in an alpine ecosystem

Understanding diet selection is important since diet determines energy intake and therefore growth of ungulate populations. Yet very few studies have reported annual variation in diet. Density-dependent diet choice by large herbivores has been reported several times, but these studies are typically either short-term or they lack replication of the density treatment. In a landscape-scale experiment with 3 replicates of two densities (25 and 80 individuals/km²) of domestic sheep, we determined diet composition using microhistological analysis during 6 summer grazing seasons (2002–2007) in alpine habitats. We tested how age class, density and temporal variation (within season, annually, and over years) affected summer diet. There was marked evidence of additive effects of these factors on overall diet composition, but interactions were few. The interaction between density and annual variation was an important determinant of the proportion of the main forage component (Avenella flexuosa), but not of the proportions of herbs, Salix spp. and for “other” forage plants. Surprisingly, the density effect on this intermediate quality forage (A. flexuosa) was not consistent among years (both positive, negative and no effects), likely arising due to large variation in the proportion of the other forage plants. We discuss how foraging ecology can supplement the insight from life history theory in explaining variation in vital rates.     

Testing for density dependence allowing for weather effects

A test for density dependence in time-series data allowing for weather effects is presented. The test is based on a discrete time autoregressive model for changes in population density with a covariate for the effects of weather. The distribution of the test statistic on the null hypothesis of density independence is obtained by parametric bootstrapping. A computer simulation exercise is used to demonstrate the gain in statistical power by allowing for weather effects. Application of the method to time-series data on three species of butterflies and two species of songbirds showed stronger evidence of density dependence than two standard tests. Received: 4 October 1996 / Accepted: 4 August 1997     

Reproductive success in reindeer males in a herd with varying sex ratio

In polygynous species, male reproductive success is often correlated with dominance status of individual males and sex ratio in the population. Reindeer, Rangifer tarandus, is a polygynous species, and here we compared the variation in male reproductive success and dominance status during two successive years in a herd with a male:female sex ratio of 1:7 and 1:3. Copulations were recorded, together with data on male dominance hierarchy and size of mating groups. Male reproductive success was estimated by paternity analysis of calves using microsatellite DNA markers. The distribution of reproductive success among the males was highly skewed for both years with the most dominant male also being the most successful. The largest mating group was established in the herd with the least skew in sex ratio. In this herd some of the adult males present were less reproductively successful than some of the more subordinate younger males. Estimates of the mating group size of males, correcting for dominance status when more than one male is present in the groups, gave good prediction of individual males' reproductive success.     

Escaping parasitism in the selfish herd: age, size and density-dependent warble fly infestation in reindeer

It has been suggested that animals may escape attack from mobile parasites by aggregating in selfish herds. A selfish herd disperses the risk of being attacked among its members and the per individual risk of parasite infection should therefore decrease with increasing animal density through the encounter–dilution effect. Moreover, in a selfish herd, dominant and agile animals should occupy the best positions and thereby receive fewer attacks compared to lower ranked animals at the periphery. We tested these predictions on reindeer ( Rangifer tarandus tarandus ) parasitized by warble flies ( Hypoderma tarandi ). Warble flies oviposit their eggs on reindeer during summer and induce strong anti-parasitic behavioural responses in the herds. In this period, reindeer are sexually segregated; females and calves form large and dense herds while males are more solitary. After hatching, the warble fly larvae migrate under the skin of their host where they encyst. In the present study encysted larvae were counted on newly slaughtered hides of male calves and 1.5 year old males from 18 different reindeer herds in Finnmark, northern Norway with large contrasts in reindeer density. In reindeer, body mass is correlated with fitness and social status and we hypothesized that individual carcass mass reflected the animal's ability to occupy the best positions within the herd. Larval abundance was higher among the 1.5 year old males than among the calves. For calves we found in accordance with the selfish herd hypothesis a negative relationship between larval abundance and animal density and between larval abundance and body mass. These relationships were absent for the 1.5 year old males. We suggest that these differences were due to different grouping behaviour where calves and females, but not males, aggregated in selfish herds where they escaped parasitism.     

Growth and condition of three introduced reindeer herds on South Georgia: the effects of diet and density

The patterns of growth and seasonal changes in body weight and fat reserves of three herds of introduced reindeer on the sub-Antarctic island of South Georgia were investigated. Two of the herds, derived from the same stock, were at different densities but the higher density herd was on the better range. In this herd there was a significant growth advantage to the male reindeer, but not to females. The third herd, of a different stock, was also at a lower density on a range which had become overgrazed. The two lower-density herds showed the same annual changes in body weight and only slight differences in the fluctuation of their fat reserves. In addition, body weights in both these herds fluctuated to the same degree as those of a barren-ground caribou in the Northern Hemisphere but fluctuations in the fat reserves of South Georgia reindeer were considerably more severe. The results suggest that the South Georgia reindeer herds are limited by forage availability.     

Patterns and strategies of winter habitat selection in alpine capercaillie

Based on radio telemetry, habitat distribution of 32 individual capercaillie Tetrao urogallus was compared with habitat availability at various levels of scale At forest stand level, capercaillie preferred large patches of late succession stages, and selected for moderate canopy closure and high abundance of feeding trees Hens and cocks selected home ranges with high proportions of old forest, and range size was related to old forest fragmentation Winter survival was high independently of habitat selection In cocks, habitat use was affected by social spacing, home range size and use of old forest were age-dependent, and habitat composition of winter ranges indirectly related to survival and mating success in spring Topography influenced habitat selection, as the birds preferred gentle slopes At the landscape scale, the distribution of bilberry Vaccinium myrtillus was the key to capercaillie habitat use, indicating that habitat requirements in autumn and spring influenced selection of winter ranges The study confirms that capercaillie in central Europe select habitats according to structural features typical of the boreal forests of the species' main distribution range However, spatio-temporal variation in resource abundance at the landscape scale sets significant limits to suitability of alpine capercaillie habitats     

Short- and long-term effects of winter and spring weather on growth and survival of red deer in Norway

Populations of red deer (Cervus elaphus) in Norway have increased continuously over the last decades. We tested the possible effects of climate and increase in population size on the survival rates and body condition of individuals in one of the northernmost populations of red deer in Europe. Based on 678 individuals of known age marked between 1977 and 1995, we estimated annual survival rates, the probabilities of being harvested and the recapture probability according to sex, age, year, winter and spring weather, population size, and, body weight and body condition, using capture-mark-recapture models. Winter harshness negatively influenced the body weight of yearlings and the survival of calves of both sexes. Spring weather influenced the survival of males in all age classes. A negative trend during the study period was detected in body weight and condition of calves and yearlings, but not in any age- or sex- specific survival rates. No significant gender differences in mean survival were shown in any age class. Moreover, there was little (male) or no (female) detectable between-year variation in survival rates for yearlings and adults. Winter weather acts as a limiting factor on population growth through a short-term effect on first-year survival and a long-term effect on body weight. We discuss the surprising low sex differences in natural survival rates and the differential effects of winter harshness on body weight, body condition and survival in relation to life history characteristics of red deer. Received: 10 November 1997 / Accepted: 2 June 1998     

Food plant selection by reindeer during winter in relation to plant quality

The relative palatabilities of nine plant taxa used as winter food by reindeer were estimated by presenting them in cafeteria and pairwise tests Feeding p reference indices were then related to the chemical composition of the plants High preference indices were obtained for the lichens Bryoria spp, Cladina arbuscula, C rangiferina and C stellaris, while low values were obtained for the moss Pleurozium schrebel and the dwarf shrubs Empetrum hermaphroditum and Vaccinium myrttllus The grass Deschampsia flexuosa and the lichen Stereocaulon paschale received intermediate scores Feeding preferences showed a strong negative correlation with acid detergent fiber (ADF)-values No statistically significant correlations were found between feeding preferences and contents of nitrogen, in vitro organic matter digestibility (IVOMD) or neutral detergent fiber (NDF)     

Rapid recovery of recently overexploited winter grazing pastures for reindeer in northern Norway

During the last 25 yr, Sami reindeer husbandry in parts of Finnmarksvidda in the Norwegian Arctic has been in a critical state because of overexploitation of lichen-dominated tundra, which serves as winter forage. To better understand the ecosystem’s capabilities for recovery we investigated vegetation cover changes over a 7-yr period, starting in 1998, at 52 sites dispersed over a large area at Finnmarksvidda. Two types of plots were established: one fenced from reindeer grazing and trampling and one open for reindeer. The investigations in 2005 showed that lichen cover had had a significant and rapid increase (up to 8.6-fold per year). The cover of vascular plants, mainly dwarf shrubs, also increased significantly, while barren areas and the cover of litter decreased significantly during the period. Mean relative growth rate of lichen biomass was 0.083 ± 0.011 per year in open plots, which is considered very rapid recovery compared to previous studies. Lichen recovery was significantly faster on leeward ridges than on exposed ridges, and fencing alone did not have any significant effects on lichen recovery, but in interaction with time, fencing contributed to increasing recovery rates. The lichen heath recovery was reciprocally correlated with reindeer density. In addition, lichen recovery was probably facilitated by recent climate changes, viz. shallower snow depths which made leeward tundra and forest floor vegetation accessible for reindeer, and increased summer precipitation rates which improved growth rates. The results from this study show that in a very short time there was a transition from an overexploited depauperate vegetation and barren ground state to a flourishing lichen-dominated vegetation state, suggesting that the injuries were repairable. The vegetation transitions which have taken place in the study area are considered to be reversible with fewer persistent effects.     

Over-winter survival in subadult European rabbits: weather effects,density dependence,and the impact of individual characteristics

The survival probability of an individual may be limited by density-dependent mechanisms and by environmental stochasticity, but can also be modified by individual characteristics. In our study, we investigated over-winter survival of subadults of an enclosed European rabbit Oryctolagus cuniculus population in a temperate zone habitat over the period 1992–2002. We: (1) selected for appropriate models to explain individual variation in over-winter survival and the animals autumn body mass, the latter was used as a measure of the individual pre-winter body condition; and (2) aimed to compare the sensitivity of the target variables on the realised variation of the factors considered. Model selection based on information theory revealed that individual over-winter survival was best explained by the combination of autumn body mass, winter temperature, population density and sex, where the probability of survival was higher in females than in males. According to this model, the probability of survival reacted most sensitively to variation in the autumn body mass and in winter temperature. Individual autumn body mass was best explained by the combination of the date of birth, population density, and weather conditions by means of the percentage of rainy days during the first 2 months after the animals had emerged above ground, where the autumn body mass was negatively related to the percentage of rainy days. The chosen model suggested that the autumn body mass reacted most sensitively to variation in the date of birth. Combining these models, we found that weather conditions during two different periods of time as well as population density, sex and the date of birth operated together to determine the probability of over-winter survival. In particular, the study points out the high impact of environmental stochasticity on over-winter survival: (1) by direct effects of winter temperature conditions, and (2) by the indirect action of weather conditions to which the animals were exposed during the early period of juvenile development.     

Coping mechanisms of alpine and arctic breeding birds: extreme weather and limitations to reproductive resilience

As ground nesting homeotherms, alpine and arctic birds mustmeet similar physiological requirements for breeding as otherbirds, but must do so in more extreme conditions. Annual springsnowfall and timing of snow melt can vary by up to 1 month anddaily temperatures near the ground surface vary from below freezingto over 45°C in alpine and arctic habitats. Species breedingin these environments have various behavioral, physiological,and morphological adaptations to cope with energetically demandingconditions. We review the ways birds cope with harsh and variableweather, and present data from long term field studies of ptarmiganto examine effects of spring weather on reproduction. In variablebut normal spring conditions, timing of breeding was not influencedby snow melt, snow depth or daily temperatures in the alpine,as breeding did not commence until conditions were generallyfavorable. Arctic ptarmigan tended to vary breeding onset inresponse to spring conditions. Generally, birds breeding inalpine and arctic habitats suffer a seasonal reproductive disadvantagecompared to birds at lower latitudes or elevations because thebreeding window is short and in late years, nest failure maybe high with little opportunity for renesting. Coping mechanismsmay only be effective below a threshold of climactic extremes.Despite strong resilience in fecundity parameters, when snowmeltis extremely delayed breeding success is greatly reduced. Alpineand arctic birds will be further challenged as they attemptto cope with anticipated increases in the frequency and severityof weather events (climate variability), as well as generalclimate warming.