Density-dependence vs. density-independence - linking reproductive allocation to population abundance and vegetation greenness

1. Recent studies have shown that optimal reproductive allocation depends on both climatic conditions and population density. We tested this hypothesis using six years of demographic data from eight reindeer (Rangifer tarandus) populations coupled with data on population abundance and vegetation greenness [measured using the Enhanced Vegetation Index (EVI)]. 2. Female spring body mass positively affected summer body mass gain, and lactating females were unable to compensate for harsh winters as efficiently as barren ones. Female spring body mass was highly sensitive to changes in population abundance and vegetation greenness and less dependent on previous autumn body mass and reproductive status. Lactating females were larger than barren females in the spring. Moreover, female autumn body mass was positively related to female autumn body mass and reproductive success and was not very sensitive to changes in vegetation greenness and population abundance. 3. Offspring autumn body mass was positively related to both maternal spring and autumn body mass, and as predicted from theory, offspring were more sensitive to changes in vegetation greenness and population abundance than adult females. A lagged cost of reproduction was present as larger females who were barren, the previous year produced larger offspring than equally sized females that successfully reproduced the previous year. 4. Reproductive success was negatively related to female autumn body mass and positively related to female spring body mass. Moreover, females who successfully reproduced the previous year experienced the highest reproductive success. The fact that negative density-dependence was only present for females that had successfully reproduced the previous year further support the hypothesis that reproduction is costly. 5. This study shows that female reindeer buffer their reproductive allocation according to expected winter conditions and that their buffering abilities were limited by population abundance and a lagged cost of reproduction and enhanced by vegetation greenness.     

Experimental evidence of cost of lactation in a low risk environment for a long-lived mammal

In a previous experiment we have documented that organisms adopt a risk-sensitive reproductive allocation when summer reproductive investment competes with survival in the coming winter ( Bårdsen et al. 2008 ). This tradeoff is present through autumn female body mass, which acts as an insurance against unpredictable winter environmental conditions. We tested this hypothesis experimentally on female reindeer experiencing stable and benign winter feeding conditions. Additional supplementary feeding and removal of newborns represented two sets of experimental manipulations. Females in the supplementary feeding group increased more in winter body mass relative to control females. This manipulation, however, did not have any effect on summer body mass development for neither females nor offspring, but we found a positive effect of feeding on offspring birth mass for smaller females. In contrast, offspring removal did have a positive effect on summer body mass development as females in this group were larger in the autumn relative to control females. In essence, we documented two immediate effects as: (1) supplementary feeding did have a positive effect on spring body mass for smaller females; and (2) offspring removal did increase the female summer somatic growth as this had a positive effect on female autumn body mass. Additionally, we tested for lagged effects, but we could not document any biologically significant effects of neither manipulation in the coming spring. The fact that we only found rather weak effects of both manipulations was as expected for risk sensitive individuals experiencing benign environmental conditions over many years.     

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.     

Reproductive success and failure: the role of winter body mass in reproductive allocation in Norwegian moose

A life history strategy that favours somatic growth over reproduction is well known for long-lived iteroparous species, especially in unpredictable environments. Risk-sensitive female reproductive allocation can be achieved by a reduced reproductive effort at conception, or the subsequent adjustment of investment during gestation or lactation in response to unexpected environmental conditions or resource availability. We investigated the relative importance of reduced investment at conception compared with later in the reproductive cycle (i.e. prenatal, perinatal or neonatal mortality) in explaining reproductive failure in two high-density moose (Alces alces) populations in southern Norway. We followed 65 multiparous, global positioning system (GPS)-collared females throughout the reproductive cycle and focused on the role of maternal nutrition during gestation in determining reproductive success using a quasi-experimental approach to manipulate winter forage availability. Pregnancy rates in early winter were normal (≥0.8) in all years while spring calving rates ranged from 0.4 to 0.83, with prenatal mortality accounting for most of the difference. Further losses over summer reduced autumn recruitment rates to 0.23–0.69, despite negligible predation. Over-winter mass loss explained variation in both spring calving and autumn recruitment success better than absolute body mass in early or late winter. Although pregnancy was related to body mass in early winter, overall reproductive success was unrelated to pre-winter body condition. We therefore concluded that reproductive success was limited by winter nutritional conditions. However, we could not determine whether the observed reproductive allocation adjustment was a bet-hedging strategy to maximise reproduction without compromising survival or whether females were simply unable to invest more resources in their offspring.     

Age-specific changes in different components of reproductive output in female reindeer: terminal allocation or senescence?

Two different processes can lead to a change in individual reproductive output with age in long-lived iteroparous vertebrates. The senescence hypothesis predicts a decline of performance in old age, whereas the terminal allocation hypothesis predicts an increase. Using long-term (>30 years) individually based data of female reindeer, we first assessed age-specific variation in body mass and different components of reproductive output. Then we investigated the contribution of senescence and terminal allocation (the increase in components of reproductive output) processes for shaping observed patterns. We found that female reindeer body mass increased up to about 11.5 years of age, and decreased afterwards, supporting the senescence hypothesis. Calf birth mass, both in absolute terms or for a given female mass, first increased and then declined with female age, also supporting the senescence hypothesis. The female mass gain (June–September) decreased with increasing age, and female change in mass between 2 consecutive years decreased with female age, all patterns again supporting the senescence hypothesis. However, the autumn calf mass did not change with age. Calf body mass in autumn tended to be positively related to female mass gain, supporting a quality effect. Raising a calf had a marked negative effect on female mass gain, indicating energetic reproductive costs of raising a calf. Calf body mass in autumn did not influence yearly female mass change. Overall, our results provided consistent evidence for general effects of senescence on most components of reproductive output and highlighted that both individual heterogeneity and reproductive costs shape female reindeer reproductive tactics.     

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.     

Pre-breeding diet affects the allocation of yolk hormones in zebra finches Taeniopygia guttata

The ability of mothers to modify offspring phenotype to match prevailing environmental conditions is an important component of reproductive success, especially in variable environments. Pre-breeding conditions, such as food abundance, may have significant consequences for both the number and quality of offspring a female produces as well as her ability to rear the offspring. In an experiment where pre-breeding diet was manipulated, we investigated if allocation of yolk androgens (testosterone and 5α-dihydrotestosterone) was related to the quality of diet experienced prior to breeding. Female zebra finches Taeniopygia guttata produced larger clutches on high quality diet than on low quality diet but with no differences in egg mass. Yolk androgen levels were repeatable within subsequent clutches of the same female and females did not change mean androgen content in eggs in relation to diet quality. However, within-in clutch pattern of yolk testosterone and DHT changed with diet treatment. Testosterone and DHT decreased with laying order on the low quality diet but remained constant on the high quality diet. Differential yolk androgen allocation within the clutch may alter the competitive differences between chicks and provide females the possibility to adjust reproductive investment and offspring phenotype already at the egg stage.     

Adaptive adjustment of offspring sex ratio and maternal reproductive effort in an iteroparous mammal

Large mammals in seasonal environments have a pattern of high-reproductive synchrony in spring, but how the timing of reproduction affects resource allocation decisions at different stages of the reproductive cycle remains largely unexplored. By manipulating the timing of conception in reindeer (Rangifer tarandus), we tested how the timing of conception affected sex ratio, gestation length and weight development of mother and offspring. Females that conceived at their first ovulation within the rut had a 60.5% probability of producing a male; in contrast, females that conceived a cycle later had a 31.3% probability of producing a male. Late conceiving females had gestation times that were 10 days shorter and the calves were 0.6 kg (9.2%) lighter at birth and 7.4 kg (14.7%) lighter in autumn. Over the year, female weight changes was similar between the groups suggesting reindeer follow a bet-hedging strategy; reducing the quality of this year's offspring to ensure their own future reproduction and survival. Harvesting is often selective leading to skewed sex ratios and age structure, which may influence the timing of reproduction due to females hesitation to mate with young males. Whenever this hesitation is strong enough to increase the frequency of recycling, harvesting is likely to have profound life history consequences.     

Role of oestradiol in the regulation of the seasonal antler cycle in female reindeer, Rangifer tarandus.

Reindeer (or caribou), Rangifer tarandus, is the only extant species of deer in which females as well as males normally develop antlers that are cast and regrown each year. This study investigated the role of ovarian oestradiol in the regulation of the seasonal antler cycle in female reindeer. Ovariectomized Norwegian reindeer living outdoors in northern Norway (69 degrees N) were treated with continuous-release subcutaneous Silastic implants containing oestradiol, which maintained the blood concentrations of oestradiol within the physiological range for the mating season from June to October-November. The treatment with oestradiol induced the synchronized maturation of the antlers and rapid cleaning of the velvet-like skin in August-September in the ovariectomized reindeer, a pattern very similar to that observed in ovary-intact controls living under the same conditions. The removal of the steroid implant in October-November caused the premature casting of the antlers in early winter in two of five animals, while the remainder cast at the normal time in spring; this response was seen whether the animals had received one or two oestradiol implants in autumn. The antlers developed by the ovariectomized, oestradiol-treated females were significantly heavier and carried more branches than the ovariectomized animals without oestradiol replacement, and were marginally heavier than the antlers of intact controls. These results support the view that oestradiol is the biologically active steroid secreted by the ovary in intact female reindeers that induces the normal development of the antlers. Oestradiol stimulates the growth and mineralization of the antler bone, the cleaning of the velvet, and suppresses the casting of the hard antlers. This endocrine control ensures that the hard antlers, which function as weapons, are retained throughout the autumn and winter when the females are normally pregnant and when competition between females over food in the snow is most intense; hence there is a reproductive advantage to explain the evolution of antlers in females.     

Senescence and costs of reproduction in the life history of a small precocial species

Species following a fast life history are expected to express fitness costs mainly as increased mortality, while slow‐lived species should suffer fertility costs. Because observational studies have limited power to disentangle intrinsic and extrinsic factors influencing senescence, we manipulated reproductive effort experimentally in the cavy (Cavia aperea) which produces extremely precocial young. We created two experimental groups: One was allowed continuous reproduction (CR) and the other intermittent reproduction (IR) by removing males at regular intervals. We predicted that the CR females should senesce (and die) earlier and produce either fewer and/or smaller, slower growing offspring per litter than those of the IR group. CR females had 16% more litters during three years than IR females. CR females increased mass and body condition more steeply and both remained higher until the experiment ended. Female survival showed no group difference. Reproductive senescence in litter size, litter mass, and reproductive effort (litter mass/maternal mass) began after about 600 days and was slightly stronger in CR than IR females. Litter size, litter mass, and offspring survival declined with maternal age and were influenced by seasonality. IR females decreased reproductive effort less during cold seasons and only at higher age than CR females. Nevertheless, offspring winter mortality was higher in IR females. Our results show small costs of reproduction despite high reproductive effort, suggesting that under ad libitum food conditions costs depend largely on internal regulation of allocation decisions.     

Heterogeneity in individual quality overrides costs of reproduction in female reindeer

Reproductive allocation at one age is predicted to reduce the probability of surviving to the next year or to lead to a decrease in future reproduction. This prediction assumes that reproduction involves fitness costs. However, few empirical studies have assessed whether such costs may vary with the age at primiparity or might be overridden by heterogeneities in individual quality. We used data from 35 years’ monitoring of individually marked semi-domestic reindeer females to investigate fitness costs of reproduction. Using multi-state statistical models, we compared age-specific survival and reproduction among four reproductive states (never reproduced, experienced non-breeders, reproduced but did not wean offspring, and reproduced and weaned offspring) and among contrasted age at primiparity. We assessed whether reproductive costs occurred, resulting in a trade-off between current reproduction and future reproduction or survival, and whether early maturation was costly or rather reflected differences in individual quality of survival and reproduction capabilities. We did not find any evidence for fitness costs of reproduction in female reindeer. We found no cost of gestation and lactation in terms of future reproduction and survival. Conversely, successful breeders had higher survival and subsequent reproductive success than experienced non-breeders and unsuccessful breeders, independently of the age at primiparity. Moreover, it was beneficial to mature earlier, especially for females that successfully weaned their first offspring. Successful females at early primiparity remained successful throughout their life, clearly supporting the existence of marked among-female differences in quality. The weaning success peaked for multiparous females and was lower for first-time breeders, indicating a positive effect of experience on reproductive performance. Our findings emphasize an overwhelming importance of individual quality and experience to account for observed variation in survival and reproductive patterns of female reindeer that override trade-offs between current reproduction and future performance, at least in the absence of harsh winters.     

Long-term shifts in the cyclicity of outbreaks of a forest-defoliating insect

Optimisation of reproductive investment is crucial for Darwinian fitness, and detailed long-term studies are especially suited to unravel reproductive allocation strategies. Allocation strategies depend on the timing of resource acquisition, the timing of resource allocation, and trade-offs between different life-history traits. A distinction can be made between capital breeders that fuel reproduction with stored resources and income breeders that use recently acquired resources. In capital breeders, but not in income breeders, energy allocation may be decoupled from energy acquisition. Here, we tested the influence of extrinsic (weather conditions) and intrinsic (female characteristics) factors during energy storage, vitellogenesis and early gestation on reproductive investment, including litter mass, litter size, offspring mass and the litter size and offspring mass trade-off. We used data from a long-term study of the viviparous lizard, Lacerta (Zootoca) vivipara. In terms of extrinsic factors, rainfall during vitellogenesis was positively correlated with litter size and mass, but temperature did not affect reproductive investment. With respect to intrinsic factors, litter size and mass were positively correlated with current body size and postpartum body condition of the previous year, but negatively with parturition date of the previous year. Offspring mass was negatively correlated with litter size, and the strength of this trade-off decreased with the degree of individual variation in resource acquisition, which confirms theoretical predictions. The combined effects of past intrinsic factors and current weather conditions suggest that common lizards combine both recently acquired and stored resources to fuel reproduction. The effect of past energy store points out a trade-off between current and future reproduction.     

Reproductive trade‐offs in a long‐lived bird species: condition‐dependent reproductive allocation maintains female survival and offspring quality

Life history theory is an essential framework to understand the evolution of reproductive allocation. It predicts that individuals of long‐lived species favour their own survival over current reproduction, leading individuals to refrain from reproducing under harsh conditions. Here we test this prediction in a long‐lived bird species, the Siberian jay Perisoreus infaustus. Long‐term data revealed that females rarely refrain from breeding, but lay smaller clutches in unfavourable years. Neither offspring body size, female survival nor offspring survival until the next year was influenced by annual condition, habitat quality, clutch size, female age or female phenotype. Given that many nests failed due to nest predation, the variance in the number of fledglings was higher than the variance in the number of eggs and female survival. An experimental challenge with a novel pathogen before egg laying largely replicated these patterns in two consecutive years with contrasting conditions. Challenged females refrained from breeding only in the unfavourable year, but no downstream effects were found in either year. Taken together, these findings demonstrate that condition‐dependent reproductive allocation may serve to maintain female survival and offspring quality, supporting patterns found in long‐lived mammals. We discuss avenues to develop life history theory concerning strategies to offset reproductive costs.     

High Female Mortality Resulting in Herd Collapse in Free-Ranging Domesticated Reindeer (Rangifer tarandus tarandus) in Sweden

Reindeer herding in Sweden is a form of pastoralism practised by the indigenous Sámi population. The economy is mainly based on meat production. Herd size is generally regulated by harvest in order not to overuse grazing ranges and keep a productive herd. Nonetheless, herd growth and room for harvest is currently small in many areas. Negative herd growth and low harvest rate were observed in one of two herds in a reindeer herding community in Central Sweden. The herds (A and B) used the same ranges from April until the autumn gathering in October–December, but were separated on different ranges over winter. Analyses of capture-recapture for 723 adult female reindeer over five years (2007–2012) revealed high annual losses (7.1% and 18.4%, for herd A and B respectively). A continuing decline in the total reindeer number in herd B demonstrated an inability to maintain the herd size in spite of a very small harvest. An estimated breakpoint for when herd size cannot be kept stable confirmed that the observed female mortality rate in herd B represented a state of herd collapse. Lower calving success in herd B compared to A indicated differences in winter foraging conditions. However, we found only minor differences in animal body condition between the herds in autumn. We found no evidence that a lower autumn body mass generally increased the risk for a female of dying from one autumn to the next. We conclude that the prime driver of the on-going collapse of herd B is not high animal density or poor body condition. Accidents or disease seem unlikely as major causes of mortality. Predation, primarily by lynx and wolverine, appears to be the most plausible reason for the high female mortality and state of collapse in the studied reindeer herding community.     

Population Densities,Vegetation Green-Up,and Plant Productivity: Impacts on Reproductive Success and Juvenile Body Mass in Reindeer

Global warming is expected to cause earlier springs and increased primary productivity in the Arctic. These changes may improve food availability for Arctic herbivores, but may also have negative effects by generating a mismatch between the surge of high quality food in the spring and the timing of reproduction. We analyzed a 10 year dataset of satellite derived measures of vegetation green-up, population densities, calf body masses and female reproductive success in 19 reindeer (Rangifer tarandus) populations in Northern Norway. An early onset of spring and high peak plant productivity had positive effects on calf autumn body masses and female reproductive success. In addition, body masses and reproductive success were both negatively related to population density. The quantity of food available, as determined by the onset of vegetation green-up and plant productivity over the summer were the main drivers of body mass growth and reproductive success. We found no evidence for an effect of the speed of spring green-up. Nor did we detect a negative mismatch between early springs and subsequent recruitment. Effects of global warming on plant productivity and onset of spring is likely to positively affect sub-Arctic reindeer.     

Age-specific variation in survival, reproductive success and offspring quality in red squirrels: evidence of senescence

Individual performance is expected to decrease with age because of senescence. We analyzed long-term data collected on a North American red squirrel population to assess the influence of age on body mass, survival and reproductive performance, and to study the effects of sex and of environmental conditions during early life on senescence patterns. Mass of males and females did not decrease at the end of life, possibly because body mass mostly reflects overall size in income breeders such as red squirrels. On the other hand, we found evidence of senescence in survival of both sexes and, to a lesser extent, in female reproductive traits. When compared to females, males had both higher survival and delayed decrease in survival, suggesting a weaker senescence in males. The offspring survival from weaning to one year of age also decreased with increasing mother age. This suggests that older females produce juveniles of lower quality, providing evidence of an intergenerational effect of mother's age on juveniles' fitness. Finally, our results indicate that variations in food conditions during early life influenced the reproductive tactics of females in the first years of their life, but not senescence patterns.     

Within-season reproductive and somatic energy allocation in a freshwater clam,Anodonta piscinalis

The effect of a change in the environment on reproductive and somatic energy allocation in an iteroparous freshwater clam Anodonta piscinalis was studied at different time of the seasonal reproductive cycle. Environmental change was produced by reciprocal transplant experiments among sites of varying productivity. In addition, clams were caged at high density to reduce the availability of resources. Transplanting females before fertilization (from May to June), or during the early development of the brood (from July to August) had no detectable effect on reproductive output. Early-season environment, however, affected body mass and percent fat content of females from two populations. This suggests that maintaining the level of reproductive allocation when resources are reduced early in the season leads to lower allocation to somatic growth and biochemical storage. Transplanting females late in the season (from September to November) had a substantial effect on reproductive output and body mass, but not on fat content, suggesting that late in the season allocation to biochemical energy storage is important. Hence, late in the season reproductive allocation may be adjusted to prevailing conditions in preparation for winter. Indeed, over-wintering site had a significant effect on percent fat content, body mass and shell growth when females were kept in a new environment from September to March. Variation among transplant sites in female body mass matched the estimated productivity of the sites, suggesting that it occurred in response to differences in the productivity of the habitats. The results emphasize the importance of taking seasonal changes in the priorities of energy allocation and the seasonality of reproductive processes into account when developing or testing models of optimal energy allocation.     

How does environmental variation influence body mass,body size,and body condition? Roe deer as a case study

We tested the influence of population density and of drought intensity (measured as the Gaussen Index in spring and summer of the year of birth) on winter body mass, hind foot length, and body condition of roe deer fawns. Body mass decreased with increasing density and increased with increasing Gaussen Index in summer, in a similar way for both males and females. Hind foot length of males showed the same response. On the other hand, hind foot length of females decreased with increasing density only after dry summers, hence when environmental conditions were very harsh. Body condition was affected neither by density nor by drought intensity. Our results indicate that body mass and size are much better indicators of phenotypic quality than body condition in roe deer. The sex-specific responses of body size to environmental conditions could correspond to a differential allocation in favour of daughters by heavier than average roe deer mothers.     

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.     

Induced orphaning reveals post-weaning maternal care in reindeer

A common by-product of human harvesting is orphaning of calves in autumn. Despite this, there are few studies in northern and temperate ungulates evaluating the fate of orphaned calves and the potential benefits to offspring and costs to mothers of post-weaning maternal care. We manipulated orphaning and forage distribution during winter for two herds of reindeer (Rangifer tarandus): one nonfed and the other supplementally fed to increase level of interference competition. Both herds consisted of females with and without calves at heel and orphaned calves. We measured survival and somatic losses during winter and distances between mother–calves and adult females–orphans within the herds. All females survived the winter, and there was no evidence of post-weaning maternal cost in terms of female’s mass loss. The winter mortality among calves was negligible and did not differ between orphans as compared to nonorphans. However, nonorphaned calves lost less mass and stayed closer to their mothers than orphans to adult females, suggesting that increased mortality might occur in harsher winters. This tended to be more marked in the fed group where interference competition was more likely due to feed being concentrated both in space and time. Reduced mass loss in nonorphans is therefore most likely due to mothers sharing and defending feeding resources and protecting their offspring from harassment by other herd members during their first winter. We conclude that hunting practise of northern and temperate ungulates where females having calf/calves at heel are intentionally or non-intentionally harvested, may have demographic side effects at least in harsh winters.