A test of alternate hypotheses for helping behavior in white-fronted bee-eaters of Kenya

Summary

Considerable controversy exists over the question of the importance of kin selection in the maintenance of helping behaviors among birds. We examined nine different hypotheses of how helpers might benefit from engaging in alloparental care activities. We break these into four categories: Through its activities the helper may A) improve its probability of surviving to the following breeding season, B) enhance its probability of becoming a breeder in the future, C) increase its reproductive success when it does become a breeder, and D) increase the production of non-descendent kin. The first three categories provide direct fitness gains to the helper; in the fourth, the benefit is indirect. The hypotheses are not mutually exclusive; rather their fitness effects are additive. Each hypothesis, however, makes specific and often separable predictions about both 1) the type of fitness benefits expected, and 2) the characteristics of the birds that serve as helpers. We tested these predictions using five year's data from a color marked and geneologically known population of white-fronted bee-eaters (Merops bullockoides) in Kenya. A) Survival was not related to status (breeder, helper, non-participant); nor did individuals living in large clans have better survival than those living in small ones. B) Newly formed pairs were equally likely to become future breeders irrespective of whether or not one or both individuals had helped previously. C) The mean number of young fledged by a first time breeder was unaffected by its prior helping experience. Neither were first time breeding pairs more likely to gain the services of others as their helpers than were pairs without prior helping experience. Taken together these results demonstrate that beeeaters gain very little direct benefit from alloparenting. D) Helpers did not enhance the survival of the breeders that they helped. But they did have a major effect in increasing nestling survival. Because bee-eater helpers are closely related to the nestlings they help to rear (average r=0.33), they obtain a large indirect benefit by increasing the production of non-descendent kin. We quantified the relative importance of indirect and direct benefits of helping (to the helper) using Vehrencamp's “kin index”, I _k (1979). I _k compares the fitness consequences of helping against an alternative strategy and calculates the proportion of the inclusive fitness gain or loss that is due to kin (indirect) benefits. Comparing the strategy sets of helping versus not helping for bee-eaters, I _k=0.89 (indicating that 89% of the benefit derived from helping is indirect). When helping was compared against the alternative of breeding, I _k=2.17. Values of I _k greater than 1.0 indicate that direct fitness gains from the alternate strategy (breeding) are greater than those from helping. The value of 2.17 indicates that the helping strategy would not be maintained except for the indirect fitness gained through the increased production of close kin. Alloparenting in white-fronted bee-eaters can thus be considered as altruistic.