Foraging competition in larger groups overrides harassment avoidance benefits in female reindeer (Rangifer tarandus).

Male harassment toward females during the breeding season may have a negative effect on their reproductive success by disturbing their foraging activity, thereby inducing somatic costs. Accordingly, it is predicted that females will choose mates based on their ability to provide protection or will aggregate into large groups to dilute per capita harassment level. Conversely, increasing group size may also lead to a decrease in foraging activity by increasing foraging competition, but this effect has rarely been considered in mating tactic studies. This study examined the importance of two non-exclusive hypotheses in explaining the variations of the female activity budget during the breeding season: the male harassment hypothesis, and the female foraging competition hypothesis. We used focal observations of female activity from known mating groups collected during the breeding season from a long-term (15 years) study on reindeer Rangifer tarandus. We found that females were more disturbed (i.e., spent less time feeding) in the presence of young dominant males, and marginally disturbed in the presence of satellite males, which supports the male harassment hypothesis. We also found that female disturbance level increased with group size, being independent of the adult sex ratio. Consequently, these results rejected the dilution effect, but strongly supported the foraging competition hypothesis. This study therefore highlights a potential conflict in female behaviour. Indeed, any gains from harassment protection were negated by an increase of 6-7 females, since adult males lead larger groups than young males.

Population density and phenotypic attributes influence the level of nematode parasitism in roe deer.

The impact of parasites on population dynamics is well documented, but less is known on how host population density affects parasite spread. This relationship is difficult to assess because of confounding effects of social structure, population density, and environmental conditions that lead to biased among-population comparisons. Here, we analyzed the infestation by two groups of nematodes (gastro-intestinal (GI) strongyles and Trichuris) in the roe deer (Capreolus capreolus) population of Trois Fontaines (France) between 1997 and 2007. During this period, we experimentally manipulated population density through changes in removals. Using measures collected on 297 individuals, we quantified the impact of density on parasite spread after taking into account possible influences of date, age, sex, body mass, and weather conditions. The prevalence and abundance of eggs of both parasites in females were positively related to roe deer density, except Trichuris in adult females. We also found a negative relationship between parasitism and body mass, and strong age and sex-dependent patterns of parasitism. Prime-age adults were less often parasitized and had lower fecal egg counts than fawns or old individuals, and males were more heavily and more often infected than females. Trichuris parasites were not affected by weather, whereas GI strongyles were less present after dry and hot summers. In the range of observed densities, the observed effect of density likely involves a variation of the exposure rate, as opposed to variation in host susceptibility.

Temporal variation in the operational sex ratio and male mating behaviours in reindeer (Rangifer tarandus).

In polygynous species, sexual selection is mostly driven by male ability to monopolize access to females in oestrous. In ungulates, the operational sex ratio (OSR), i.e. the proportion of males to individuals ready to mate, varies throughout the peak rut, resulting from the temporal variation in the number of females in oestrous. But the way males adjust their mating tactics to maximise their access to females in oestrous (i.e. as OSR varies) is yet to be investigated. Using 15 years of behavioural observations in reindeer (Rangifer tarandus), we compared the relative importance of time within the rutting season (days to the peak-rut) and the OSR to explain the variation in the propensity (i.e. the frequency after controlling for the potential number of encounters) of young and adult dominant males to engage in four mating tactics: herding females, chasing other males, investigating female reproductive status, and courting females. Male-male agonistic behaviour was the most frequent mating behaviour, followed by herding. As predicted, dominant male mating tactics changed over the rutting season: first herding, then chasing other males, and finally investigating and courting females. In contrast to our prediction, we did not find support for the OSR theory. We noted some discrepancies in how young and adult dominant males adjusted their tactics during the mating season, adults being more efficient in timing and in performing their behaviour to maximise access to females in oestrous. The reported sequence of mating tactics may be more efficient than a static mating tactic to monopolize females in oestrous, regardless of the population composition.

Highly competitive reindeer males control female behavior during the rut.

During the rut, female ungulates move among harems or territories, either to sample mates or to avoid harassment. Females may be herded by a male, may stay with a preferred male, or aggregate near a dominant male to avoid harassment from other males. In fission-fusion group dynamics, female movement is best described by the group's fission probability, instead of inter-harem movement. In this study, we tested whether male herding ability, female mate choice or harassment avoidance influence fission probability. We recorded group dynamics in a herd of reindeer Rangifer tarandus equipped with GPS collars with activity sensors. We found no evidence that the harassment level in the group affected fission probability, or that females sought high rank (i.e. highly competitive and hence successful) males. However, the behavior of high ranked males decreased fission probability. Male herding activity was synchronous with the decrease of fission probability observed during the rut. We concluded that male herding behavior stabilized groups, thereby increasing average group size and consequently the opportunity for sexual selection.