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.     

Wear Fast,Die Young: More Worn Teeth and Shorter Lives in Iberian Compared to Scottish Red Deer

Teeth in Cervidae are permanent structures that are not replaceable or repairable; consequently their rate of wear, due to the grinding effect of food and dental attrition, affects their duration and can determine an animal''s lifespan. Tooth wear is also a useful indicator of accumulative life energy investment in intake and mastication and their interactions with diet. Little is known regarding how natural and sexual selection operate on dental structures within a species in contrasting environments and how these relate to life history traits to explain differences in population rates of tooth wear and longevity. We hypothesised that populations under harsh environmental conditions should be selected for more hypsodont teeth while sexual selection may maintain similar sex differences within different populations. We investigated the patterns of tooth wear in males and females of Iberian red deer (Cervus elaphus hispanicus) in Southern Spain and Scottish red deer (C. e. scoticus) across Scotland, that occur in very different environments, using 10343 samples from legal hunting activities. We found higher rates of both incisor and molar wear in the Spanish compared to Scottish populations. However, Scottish red deer had larger incisors at emergence than Iberian red deer, whilst molars emerged at a similar size in both populations and sexes. Iberian and Scottish males had earlier tooth depletion than females, in support of a similar sexual selection process in both populations. However, whilst average lifespan for Iberian males was 4 years shorter than that for Iberian females and Scottish males, Scottish males only showed a reduction of 1 year in average lifespan with respect to Scottish females. More worn molars were associated with larger mandibles in both populations, suggesting that higher intake and/or greater investment in food comminution may have favoured increased body growth, before later loss of tooth efficiency due to severe wear. These results illustrate how independent selection in both subspecies, that diverged 11,700 years BP, has resulted in the evolution of different longevity, although sexual selection has maintained a similar pattern of relative sex differences in tooth depletion. This study opens interesting questions on optimal allocation in life history trade-offs and the independent evolution of allopatric populations.     

Winter grazing of reindeer in woodland lichen pasture: Effect of lichen availability on the condition of reindeer

Winter grazing of semi-domesticated reindeer (Rangifer t. tarandus) was investigated at the woodland lichen pasture (lichen approximately 550 kg DM ha⁻¹) in Kaamanen, northern Finland during the winter 1996–1997. Nine female reindeer mainly dug their food in the snow for 122 days (3 December–4 April) in a fenced area of 36.3 ha. Over half of the fenced area was lichen dominated dry pine forest. The amount of lichens in lichen forest inside the fence was estimated before and after grazing. Area of grazed and condition of reindeer as well as snow conditions were monitored. Reindeer grazed over the whole area of lichen forest in early winter but from mid-winter they tended to graze on the areas with the greatest lichen abundancy. The amount of lichens measured decreased in the latter areas by 40% and in the other part of the lichen pasture by 17%, respectively. In both of these areas the residual amounts of lichens left after grazing were similar. Of the dominant lichens, the amount of C. stellaris decreased the most and the amount of Cl. uncialis the least. During the study, the estimated average daily area grazed varied from 4 to 87 m² per reindeer. It was calculated that individual reindeer obtained 2.6 kg of lichen DM per day during the most intensive digging period when the body condition score and weight of reindeer increased. Otherwise, the body condition score and weight decreased. Reindeer finished foraging for ground lichens and started to search for arboreal lichens in mid-March when the snow layer was 70–80 cm thick and contained some hard snow layers which lifted reindeer. Both the amount of lichens in the pasture and the snow conditions essentially affect the nutritional status of reindeer in the woodland region during winter. Assuming that a reindeer is able to graze around 30 m² per day in the snow during mid and late winter, there should be, on the basis of energy demand and grazing behaviour of reindeer as well as the nutritive value of lichen, an estimated 1000 kg lichen DM ha⁻¹ available in a good condition woodland lichen pasture.     

Nutritive value of terricolous lichens for reindeer in winter

Abstract:In contrast to virtually all other species of ruminants, reindeer (Rangifer tarandus) eat substantial amounts of lichens in winter. Several different species of lichens are eaten and most are highly palatable. The chemical composition and the in vitro digestibility of six species of terricolous lichens commonly eaten by reindeer were measured. Ruminal inoculum was obtained in winter from six free-ranging reindeer that had been grazing on natural pastures in northern Norway. Analysis of the plant parts recovered from their rumens confirmed that the animals had been eating a mixed diet that included both lichens and vascular plants. The chemical composition and the in vitro digestibility of the different species of lichens varied considerably between species and genera. Cetraria islandica, C. nivalis andCladonia arbuscula were highly digestible [69–77% dry matter (DM)], Stereocaulon paschale was poorly digestible (43% DM), whereas Cladonia stellaris and C. gracilis had an intermediate digestibility (56–57% DM). Mixing Cladonia stellaris with vascular plants (50:50) had no effect on the in vitro digestibility of the combined substrates. The in vitro digestibility of Cladonia stellaris in inoculum from two captive reindeer which had had no access to lichens was very low (10% DM). This shows that the source of inoculum used for digestibility trials has a major effect on the apparent digestibility of the substrates. Evidently, the extent to which reindeer are able to utilize lichens depends on the species that are selected and on what the animals have been eating recently.     

Risk‐sensitive reproductive allocation: fitness consequences of body mass losses in two contrasting environments

For long‐lived organisms, the fitness value of survival is greater than that of current reproduction. Asymmetric fitness rewards suggest that organisms inhabiting unpredictable environments should adopt a risk‐sensitive life history, predicting that it is adaptive to allocate resources to increase their own body reserves at the expense of reproduction. We tested this using data from reindeer populations inhabiting contrasting environments and using winter body mass development as a proxy for the combined effect of winter severity and density dependence. Individuals in good and harsh environments responded similarly: Females who lost large amounts of winter body mass gained more body mass the coming summer compared with females losing less mass during winter. Additionally, females experienced a cost of reproduction: On average, barren females gained more body mass than lactating females. Winter body mass development positively affected both the females' reproductive success and offspring body mass. Finally, we discuss the relevance of our findings with respect to scenarios for future climate change.     

Winter climate change: Ice encapsulation at mild subfreezing temperatures kills freeze-tolerant lichens

While it has been widely proven that many lichens are extremely freeze-tolerant in the dry state, little is known about how moist lichens respond to freezing under oxic and anoxic conditions. In circumpolar areas where lichens are an important component of boreal and Arctic ecosystems, winter climate is changing, leading to increased frequency of winter thaw and ground-icing events. It is imperative to elucidate in further detail how northern vegetation responds to being encapsulated in ice. A winter icing simulation experiment was therefore undertaken, encapsulating two reindeer lichens (Cladonia stellaris and Cladonia rangiferina) and two epiphytic lichens (Parmeliopsis ambigua and Melanohalea olivacea) in ice at temperatures just below freezing for a maximum period of 98 d. Photosynthetic and chlorophyll fluorescence rates decreased strongly, clearly suggesting that the algal partner of the lichens was dying. Low but detectable respiration rates indicate that the fungal partner maintained some physiological activity. Ethanol accumulated in the lichens during ice encapsulation as a result of anaerobic respiration. The algae probably were dying from a combination of depletion of carbon reserves and toxic levels of ethanol and other stress metabolites. This experiment shows that boreal and Arctic-alpine lichens are sensitive to a warmer and more fluctuating winter climate. Increasing frequency of winter icing events may therefore have extensive and hitherto unknown effects on lichen-dominated ecosystems, their herbivores and the indigenous peoples whose livelihoods largely depend on them.     

Adaptive regulation of body reserves in reindeer, Rangifer tarandus: a feeding experiment

The costs and benefits of body reserves fluctuate according to predictable factors such as season and life-cycle stage. Theory suggests that individuals at any time should regulate their body reserves according to the current balance between costs and benefits. Most studies on adaptive body mass regulation have been done on small passerine birds. In large vertebrates the costs associated with body reserves are assumed to be small and the reserves of these species are therefore thought to be dictated by environmental limitations. In this study we present experimental evidence for adaptive body mass regulation in female semi-domesticated reindeer ( Rangifer t. tarandus ). The risk of starvation in this species is highest in late winter. During snow melt this risk is reduced and the females should direct their effort towards the protection of their new born calf. To test how these seasonal and life-cycle changes are related to body mass regulation, we conducted a crossed experiment with two treatments where females were fed ad libitum during winter and spring respectively. During winter, the females from the fed group gained on average 12% of their initial body weight while the females from natural pastures lost on average 6% of their initial body weight. This strong response to winter feeding had no effect on reproductive performance, and the previously fed females lost their excess of body reserves during feeding in spring. This suggests that body reserves during winter primarily is used as an insurance against stochastic periods of starvation and that the females regulate their body reserves down to a set point in spring when the risk is reduced. We found however a positive correlation between initial female body weight and reproductive performance suggesting a close relationship between body weight and intrinsic individual qualities.     

Ice encapsulation protects rather than disturbs the freezing lichen

Arctic and alpine terricolous lichens are adapted to harsh environments and are tolerant to extremely low temperatures when metabolically inactive. However, there are reports indicating that freezing can be lethal to metabolically active lichens. With a projected warmer and more unstable climate, winter precipitation at high latitudes will fall more frequently as rain, causing snowmelt and encapsulating terricolous lichens in ice or exposing them to large temperature fluctuations. Lichens are a major winter food source for reindeer in most parts of the circumpolar region. A laboratory experiment tested how three hydrated reindeer forage lichen species covered by snow, encapsulated in ice, or uncovered responded to storage at freezing temperatures and subsequent warming. Photosynthetic performance (maximal fluorescence of dark-adapted samples and net photosynthetic rates) was significantly lower in lichens not insulated by snow or ice, whereas there were few differences between the snow and ice treatments. It is suggested that snow and ice provide sufficiently moist environments to improve extracellular and reduce intracellular ice nucleation activity. Ice encapsulation, which is often lethal to vascular plants, did not have any negative effects on the studied lichens. The results indicate that complete snow and ice melt followed by refreezing can be detrimental to terricolous lichen ecosystems. Reduced lichen biomass will have a negative effect both on reindeer winter survival and the indigenous peoples who herd reindeer.     

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.     

Evolutionary responses to a changing climate: Implications for reindeer population viability

If we want to understand how climate change affects long‐lived organisms, we must know how individuals allocate resources between current reproduction and survival. This trade‐off is affected by expected environmental conditions, but the extent to which density independent (DI) and density dependent (DD) processes interact in shaping individual life histories is less clear. Female reindeer (or caribou: Rangifer tarandus) are a monotocous large herbivore with a circumpolar distribution. Individuals that experience unpredictable and potentially harsh winters typically adopt risk averse strategies where they allocate more resources to building own body reserves during summer and less to reproduction. Such a strategy implies that the females do not reproduce or that they produce fewer or smaller offspring. A risk averse strategy thus results in females with large autumn body reserves, which is known to increase their survival probabilities if the coming winter is harsh. In contrast, females experiencing predictable winters may adopt a more risk prone strategy in which they allocate more resources to reproduction as they do not need as many resources to buffer potentially adverse winter conditions. This study uses a seasonal state‐dependent model showing that DD and DI processes interact to affect the evolution of reproductive strategies and population dynamics for reindeer. The model was run across a wide range of different winter climatic scenarios: One set of simulations where the average and variability of the environment was manipulated and one set where the frequency of good and poor winters increased. Both reproductive allocation and population dynamics of reindeer were affected by a combination of DI and DD processes even though they were confounded (harsh climates resulted in lowered density). Individual strategies responded, in line with a risk sensitive reproductive allocation, to climatic conditions and in a similar fashion across the two climatic manipulations.     

Allocating protein to reproduction in arctic reindeer and caribou

Reindeer (Rangifer tarandus tarandus) and caribou (Rangifer tarandus granti) use body stores (capital) and food intake (income) for survival and reproduction. Intakes of low-nitrogen (N) food declined in winter and increased in spring (51-83 g dry matter kg(-0.75) d(-1)). Reindeer calved before regaining food intake, whereas caribou calved 28 d later. Body N was conserved by minimizing oxidation of amino acid N to urea. Maternal protein stored from early winter was used for 96% of fetal growth in reindeer but only 84% of fetal growth in later-birthing caribou. Both subspecies rely on maternal body protein for 91% of the protein deposited in the neonate via milk over the first 4 wk. All females lost body protein over winter, but lactating females continued to lose protein while nonreproductive females regained protein. Net costs of lactation above maintenance were greater for N (110%-130%) than for energy (40%-59%). Large fat stores in reindeer spare body protein from oxidation in winter, whereas in caribou, less fat with the same body protein favors migration when food is inadequate. The resilience of Rangifer populations to variable patterns of food supply and metabolic demand may be related to their ability to alter the timing and allocation of body protein to reproduction.     

Sex-specific strategies of dentine depletion in red deer

Worn teeth in herbivore ungulates may be related to lower efficiency in mastication and hence lower performance. However, selection should favour maximal performance in terms of body mass and reproductive capacity during reproductive lifespan, when permanent teeth are already partially worn. We hypothesize that wear rate may respond to a strategy of use of tooth materials (notably dentine), which balances instantaneous wear rate and performance against tooth preservation for future performance and reproduction. In the present study, we investigated 4151 carcasses of Iberian red deer Cervus elaphus hispanicus and show that more worn molars were not related to lower performance throughout age. By comparing between sexes, tooth wear rates were smaller in females than in males, but the relationship between tooth wear and body performance also differed between the sexes: females did not show a significant relationship between tooth wear and performance but males with more worn teeth were in general heavier and had larger antlers until senile age, when more depleted teeth were related to smaller antlers. These results reveal, for the first time, sex-specific lifetime strategies of dentine expenditure: maintenance of performance ability throughout a longer reproductive lifespan in females, compared with maximizing current performance by depleting dentine reserves within a shorter lifespan in males.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 487–497.     

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.     

Fitness consequences of alternative life histories in water striders,Aquarius remigis (Heteroptera: Gerridae)

Using field and laboratory observations and experiments over 3 years, I investigated whether reproductive trade-offs shape individual life histories in two natural populations of the water strider, Aquarius remigis, in which univoltine and bivoltine life cycles coexist. Both later eclosion dates and food shortages, even after adult eclosion, induced diapause in females, thus deferring reproduction to the following spring. Adult body size was positively affected by food availability during juvenile development. Higher food levels also increased the reproductive output of females, but not their longevity or oviposition period. When compared to spring breeders (univoltine life cycle), direct (summer) breeders (bivoltine life cycle) experienced reduced lifetime egg numbers and longevity, as well as reduced survivorship of their second-summer-generation offspring; these reproductive costs offset, at least in part, the advantage in non-decreasing populations of having two generations per year. Fecundity was correlated with body size, and among summer-generation females direct breeders were larger than non-breeders. The time remaining before the onset of winter and/or the time since adult eclosion augmented cumulative energy uptake, and consequently the lipid reserves and winter survival probability of non-breeding (diapausing) summer adults approaching hibernation. Overwintered spring reproductives died at faster rates than non-reproductive summer individuals despite greater food availability in spring, indicating a mortality cost of reproduction. Body length correlated with absolute and not with proportional lipid content but showed no consistent relationship with survivorship in the field. These results are in agreement with current theory on the evolution of insect voltinism patterns, and further indicate high degrees of life history flexibility (phenotypic plasticity) in the study populations in response to variable environmental factors (notably photoperiod and food availability). This may be related to their location in a geographic transition zone from uni- to bivoltine life cycles.     

Functional Anatomy of the Omasum in High Arctic Svalbard Reindeer (Rangifer tarandus platyrhynchus) and Norwegian Reindeer (Rangifer tarandus tarandus)

The structure and fill of the omasum was investigated in summer and in winter in adult female reindeer living on the polar desert and tundra of the high Arctic archipelago of Svalbard and in sub-Arctic mountain habitats in northern Norway The mean total mass of the omasum in non-lactating adult female Svalbard reindeer was 467 g (0.65 g per 100 g live body mass (BM)) in September and 477 g (1.03 g per 100 g BM) in April. By contrast, the mean mass of the omasum in non-lactating adult reindeer in northern Norway was 534 g (0.83 g per 100 g BM) in September but only 205 g (0.35 g per 100 g BM p<0.05) in late March, owing to a decrease in both tissue mass and the wet mass of the contents of the organ. The mean absorptive surface of the omasum in Svalbard reindeer was 2300 cm2 in September and 2023 cm2 in April. In Norwegian reindeer, by contrast, the absorptive surface area decreased from 2201 cm2 in September to 1181 cm2 (p<0.05) in late March. The marked seasonal decline of omasal tissue and contents in Norwegian reindeer probably results from intake of highly digestible forage plants, including lichens, in winter. Svalbard reindeer, a non-migratory sub-species, survive eating poor quality fibrous vascular plants in winter. The absence of any marked seasonal change in the mass, total absorptive surface area or filling of the omasum in Svalbard reindeer in winter despite a substantial decline in body mass presumably reflects their need to maintain maximum absorption of nutrients, including volatile fatty acids, when feeding on such poorly fermentable forage.     

Trapping oestrid parasites of reindeer: the relative age, fat body content and gonotrophic conditions of Cephenemyia trompeand Hypoderma tarandifemales caught in baited traps

Abstract. Dissection of flies caught in northern Norway revealed that only mated, gravid females of H.tarandi (L.) and larviparous C.trompe (Modeer) were caught in host-mimicking C0₂-baited traps. Trapped females had the same gonotrophic and fat body (FB) conditions as females caught on and around reindeer. Most trapped females of both species were of middle to old age, having only one-half to no FB reserves left and only a few to moderate numbers of eggs or larvae remaining. Most young females trapped also had previously oviposited or larviposited at least once, and some newly eclosed, mated females were present throughout the fly season. Based on the known location of reindeer herds, it was evident that trapped flies that had recently oviposited or larviposited had dispersed into the trapping area from 25–100 km away. Declining FB reserves sustained wild-caught C.trompe females (and in utero larvae) in the laboratory for 14 days and H.tarandi females for 18 days. Reserve FB also was depleted during long flights. Females of both species that flew for the longest times (5.1–11.7 h) on a laboratory flight mill had low FB reserves, but nearly maximum numbers of eggs or larvae. Conversely, most females that flew for less than 5 h on the flight mill had little or no FB remaining, and few eggs or larvae. The large FB reserves accumulated as larvae feed in the vertebrate host enable the non-feeding adults to survive and infect their hosts even after prolonged periods of flight-inhibiting climatic conditions.     

Observational evidence of risk-sensitive reproductive allocation in a long-lived mammal

Organisms should adopt a risk-sensitive reproductive allocation when summer reproductive allocation competes with survival in the coming winter. This trade off is shown through autumn female body mass, which acts as an insurance against unpredictable winter environmental conditions. We tested this hypothesis on female reindeer in a population that has experienced a time period of dramatic increase in abundance. Environmental conditions during winter were fairly stable (with the exception of 1 year). We conclude that increased population abundance (perhaps in interaction with winter environmental conditions) could have represented a worsening of winter environmental conditions as both autumn offspring and spring female body mass decreased during the course of the study. Moreover, we found that the cost of reproduction was related to environmental conditions as: (1) autumn body mass was larger for barren than for lactating females, and this difference was temporally highly variable; (2) lactating females produced smaller offspring than barren ones in the following year; and (3) reproductive output (offspring size) decreased over time. We also found evidence of quality effects as lactating females had a higher reproductive success in the following year. In sum, a worsening of winter conditions lead to: (1) decreased reproductive output; (2) lowered autumn body mass for lactating females; and (3) increased body mass for barren females. Since females reduce their reproductive allocation as winter conditions becomes more severe, we conclude that reindeer have adopted a risk-sensitive reproductive allocation.     

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.     

Winter habitat–space use in a large arctic herbivore facing contrasting forage abundance

We analysed how changes in resource levels influence foraging trade-offs in late winter by wild Svalbard reindeer. Forage plants, and particularly lichens, were less abundant at the overgrazed Brøggerhalvøya compared with the neighbouring Sarsøyra. Strong interactions occurred between habitat selection, home range size, and feeding crater selection. At Brøggerhalvøya, radiocollared females generally selected productive habitat (high summer NDVI; Normalised Difference Vegetation Index). “Immigrants” at Sarsøyra (dispersed from Brøggerhalvøya in early winter) had similar habitat preferences, probably due to past experience. In contrast, “residents” at Sarsøyra were more influenced by abiotic conditions, using habitat with low NDVI, but selecting for high-quality forage (lichens) when cratering. This suggests more quality-based selection at the expense of quantity when forage abundance increases. Habitat–space use relationships also differed between the animal categories, as home range size decreased with availability of preferred habitat. Thus, changes in forage abundance can strongly influence winter habitat–space use interactions in predator-free systems.     

Seasonal grazing effects by semi-domesticated reindeer on subarctic mountain birch forests

In northern Fennoscandia, the spatial and temporal grazing practices of semi-domesticated reindeer (Rangifer tarandus tarandus) vary, which implies different grazing effects dependent on natural conditions as well as management regime (i.e., timing and intensity of grazing). We compared density and biomass of main plant groups in semi-dry mountain birch forests exposed to either long-term summer or winter grazing in three reindeer herding districts in the northernmost Finland. Percent plant cover, height, and biomass of reindeer lichens (Cladonia spp.) and dwarf shrubs (Vaccinium uliginosum, Calluna vulgaris, and Betula nana) were lower on summer ranges compared with winter ranges. The biomass of other dwarf shrubs (Vaccinium myrtillus and V. vitis-idaea), and graminoids and herbs, and the % cover of non-vegetated bare soil and litter were, however, higher on summer ranges than on winter ranges. Young mountain birch shoots (Betula pubescens ssp. czerepanovii) were less frequent on summer ranges than on winter ranges. The total leaf biomass under the browsing height of reindeer (<1.5 m) was also lower on summer ranges compared with winter ranges. Especially in drier and nutrient poor mountain birch forests, intensive summer grazing reduces the quantity of lichens and total plant biomass which affects the ecological state and productivity of these forests and also reduces especially their winter grazing value for reindeer. Therefore, in addition to regulating the maximum sustained numbers of reindeer, pasture rotation systems that effectively protect dry and nutrient poor vegetation from summer grazing and trampling should be encouraged.