Fitness consequences of helping behavior in the western bluebird

Abstract

We examined the fitness consequences of helping behavior in the western bluebird (Sialia mexicand) at Hastings Reservation in Carmel Valley, California, USA, and tested hypodieses for how helpers benefit from engaging in alloparental behavior. Both juvenile and adult western bluebirds occasionally help at the nest. During a 12 year period, all adult helpers and most juvenile helpers were male. Helpers usually fed at nests of both their parents and rarely helped when only one parent was present. The frequency of pairs with adult helpers was only 7%, but nearly one-third of adult males helped among those with both parents on the study area. At least 28% were breeders whose nests failed. The propensity to help appears to depend upon parental survival, male philopatry, and die breeding success of potential helpers. Feeding rates were not increased at nests with juvenile helpers, apparendy because breeding males reduced their feeding rates. In contrast, adult helpers increased die overall rates of food delivery to die nest in spite of a reduction in die number of feeding trips made by both male and female parents. Helpers did not derive any obvious direct fitness benefits from helping, but diey had greater indirect fitness than nonhelpers due to increases in nesding growdi rates and fledging success at dieir parents' nests. Helpers fledged fewer offspring in their first nests dian did nonhelpers, suggesting diat they were birds widi reduced reproductive potential. Aldiough we have not yet measured the effect of extrapair fertilizations on die fitness benefits of helping, we calculated die difference in fitness between helpers and nonhelpers as a function of die potential helper's paternity when breeding independendy and his fadier's paternity in die nest at which he might help. In conjunction with constraints on breeding and indirect fitness benefits, we predict that relatedness of males to die young in dieir own as well as dieir parents' nests will influence helping behavior in western bluebirds.