Within-group relatedness and patterns of reproductive sharing and cooperation in the tropical chestnut-crested yuhina

In cooperatively breeding animals, genetic relatedness among group members often determines the extent of reproductive sharing, cooperation and competition within a group. Studies of species for which cooperative behaviour is not entirely based on kinship are key for understanding the benefits favouring the evolution and maintenance of cooperative breeding among nonrelatives. In the cooperatively breeding chestnut-crested yuhina, Yuhina everetti, a songbird endemic to Borneo, we tested whether unrelated helpers are more likely to gain parentage than are related helpers consistent with the hypothesis that inbreeding risk constrains reproduction by related helpers. We also examined whether related or unrelated helpers provision broods more because of differences in their potential indirect or direct fitness benefits of helping. Kin structure of breeding groups (breeding pair and up to eight helpers of both sexes, median = 2 helpers, 96% of 57 pairs had helpers) based on genetic analysis was mixed; 48% of 76 breeder/helper dyads were first-order (26%) or second-order (22%) relatives of one or both members of the breeding pair, and 52% were nonrelatives. Only unrelated male and female helpers gained parentage, and helpers did not differ in their provisioning rate according to their relatedness to the broods. We documented quasi-parasitism or co-breeding by female helpers in 14% of 29 broods and extrapair paternity by male helpers in 21% of 47 broods. This rate of extrapair paternity is relatively high among the few tropical species examined but fit with predictions for mixed-kin groups where inbreeding is avoided. These findings support the emerging pattern for cooperative breeding in birds with mixed-kin groups, wherein unrelated helpers are more likely to gain parentage than are related helpers and helping effort is not necessarily predicted by kinship.