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The consequence of helping behavior on breeders fitness is still controversial. We used multivariate analyses to investigate for the effects of male and female subordinates on breeders’ components of fitness in the Alpine marmot, Marmota marmota. We found that male and female subordinates, respectively, increased and decreased juvenile survival during winter. Thus, we give evidence that male subordinates should be considered as helpers, and that helpers provided breeders with immediate reproductive success gains, whereas subordinates females were costly. Helpers had no positive effects on female body condition, on persistence (future survival) of dominants, and on future reproduction (occurrence and size of a litter). Helpers thus did not provide breeders with delayed fitness benefits, and therfore, the load-lightening hypothesis was not supported. On the contrary, helpers had delayed fitness cost for dominant males and, consequently, for dominant females. Immediate benefits counterbalanced by delayed costs suggested an optimal number of helpers in the family group both from male and female perspectives. An optimality model well predicted the observed mean number of helpers in Alpine marmot family groups. Optimal numbers of helpers were slightly different for males and females, suggesting a potential conflict of interest between dominants. We finally discuss the possible mechanisms of helping that may explain the observed pattern in the Alpine marmot. Key words: cooperative breeding, loadlightening hypothesis, Marmota marmota, mixed models, optimality. [Behav Ecol 15:916–924 (2004)]

In cooperatively breeding vertebrate species, a clear theoretical prediction about the direction of sex ratio adjustment can be made: mothers should bias the sex ratio of their offspring towards the helping sex when helpers are absent. A consistent trend in the direction predicted by theory exists in cooperative birds, but theory is still poorly tested in cooperative mammals. Here, multivariate analyses are applied to a long-term data set to test this prediction in two ways in the alpine marmot: (1) across females in a population and (2) in individual females across multiple years. It was shown that in the alpine marmot offspring sex ratio was biased towards the helping sex (males) when helpers were absent, whereas helped mothers produced unbiased sex ratio. Unhelped mothers did not adjust the litter size but produced more sons and fewer daughters than helped mothers. These results support the theoretical prediction and explain well the male bias observed among juvenile alpine marmots at the population level. The occurrence of possible sex ratio manipulations in cooperatively breeding vertebrates is also discussed. Copyright 2004.

In a French population of Alpine marmots (Marmota marmota), the sex ratio at weaning was biased in favor of males. This bias also seemed to exist at birth. Under Fisher’s equal allocation principle, this means that daughters should be more costly to produce than sons. Because the Alpine marmot can be considered a cooperative breeding species, we investigated whether the differential cost between sons and daughters may be explained by the helper repayment hypothesis. The Alpine marmot uses social thermoregulation during hibernation, allowing juveniles to better survive over winter. In the study population, juvenile survival during winter increased with group size. More precisely, juvenile survival during winter increased with the number and with the proportion of subordinate males in the hibernating group, but juvenile survival did not depend on the number of subordinate females. As our results did not support alternative hypotheses to explain the observed bias in sex ratio among offspring at emergence, we conclude that the helper repayment hypothesis is the best candidate to explain the observed offspring sex ratio bias in Alpine marmots. By participating in social thermoregulation, subordinate males may repay part of the investment they received from their parents and thus become less costly to produce. We suggest that only subordinate males helped because they may gain direct fitness benefits, whereas subordinate females may only expect indirect fitness benefits from helping. Finally, the offspring sex ratio per individual parent was male biased, but mothers adjusted the size and the sex composition of their litters according to their phenotypic condition as expected from the Trivers-Willard hypothesis. Key words: Alpine marmots, cooperative breeding, helper repayment hypothesis, Marmota marmota, sex ratios. [Behav Ecol 11:507–514 (2000)]