In sexually dimorphic ungulates, male reproductive success depends on fighting with other males for access to females during a brief rutting season. Large body size is necessary for success in intrasexual competition, and a few large-sized males are often able to monopolize access to female groups. Earlier studies have reported that reproductive effort increases with age until prime-age is reached, and one study that population density lowered effort in (older) males. No study has directly assessed whether there is within-age-class variation in effort resulting from varying levels of intra-male competition. It is reported here the weight loss during the rutting season of 54 individual male reindeer Rangifer tarandus coming from eight herds with varying density (3.3-6.0 deer/km 2 ) and sex ratio (4-28% males). In agreement with earlier studies, reproductive effort was lower for young (1-to 2-year-old) than for prime-aged (3-to 5-year-old) males both on an absolute and relative scale. Among 1-year-old males (n = 33), effort was lower as sex ratio became closer to even, but density during the rutting season had no effect. This suggests that yearling males take a more active role when prime-aged males are absent. In addition to the insight into male ungulate life history, understanding male rutting behaviour may also have implications for population dynamics.

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.

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.

In polygynous mammals, fitness differences may reflect differences in phenotypic quality as well as experience. This study determines dominance hierarchy among female reindeer Rangifer tarandus from two experimental herds (consisting of c. 45 animals in each) during 2 consecutive years. The influence of body mass, antler size and age on social rank in the herds was investigated, first using simple regression analysis. The combined effect of body mass, age and antler size on female rank was further assessed using principal component analysis, as these three parameters were significantly correlated. The improved Laundau linearity index of c. 0.5 (P r < 0.001) in both herds indicated that a substantial part of the hierarchies was explained by their linearity properties. Consistently, body mass, female age and antler size, as well as their combined effect (measured by the scores on the main axis, PC1), influenced social rank in all four groups. It was concluded that both body mass and age are good predictors of social rank in female reindeer, whereas antler size in comparison plays a less important role in herds with a 'normal' female age structure. This suggests that female antlers may have evolved in intersexual rather than intrasexual competition. The temporal variations in the importance of body mass and age, probably owing to variation in female age structure between the 2 years, calls for conservative interpretations of whether body mass or age is more important in determining social rank among female reindeer. This is confirmed by the PCA analyses, where all three variables contributed more or less equally to the first component, the size variable, which on average explained more of the variation in female rank than body mass and age, suggesting that phenotypic quality expressed as the combination of the three variables is a better predictor of social rank than the variables per se. Hence, general conclusions about social rank based on single studies including few animals may not be credible.

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