Experimental tests of endocrine function in breeding and nonbreeding raptors

Many long-lived avian species defer reproduction for several years, often displaying a "floating" behavior characterized by the lack of mates and exclusive territories. Understanding the proximate mechanisms regulating floating behavior is a relevant topic of research for physiologists, behavioral ecologists, and population biologists because a prolonged period of nonbreeding can negatively affect lifetime fitness and change population dynamics. Here we tested two hypotheses linking endocrine function to floating status: (a) floaters undergo a period of sexual immaturity characterized by lower gonadal function (hypothesis of sexual immaturity), and (b) floating status is socially imposed by dominant conspecifics and revealed by the adrenocortical response to stress (hypothesis of social subordination). The two hypotheses were tested in a population of free-living black kites Milvus migrans in Doñana National Park (southwest Spain), where breeders coexist with young floaters that defer reproduction for 3-7 yr. Hypophysial-gonadal function, estimated as androgen production in response to experimental challenge with gonadotropin-releasing hormone (c-GnRH-I), was similar in magnitude and timing between floating and breeding males. The same treatment was, however, unable to elicit any response in terms of increasing estradiol or total androgen levels in females regardless of their breeding status. Following experimental capture and restraint, the adrenocortical response to stress (estimated as circulating corticosterone levels) was higher in floating than in breeding males, while females showed the opposite pattern (i.e., lower response to stress in young floaters compared with breeders). Contrary to the hypothesis of sexual immaturity, our results suggest that floating males are physiologically capable of reproducing. The reported differences in adrenocortical function support the idea that floaters are socially subordinate to breeders, and corticosterone responses reflect the sex-specific roles during competition in socially monogamous species.