Individual differences in protandry, sexual selection, and fitness

Protandry is the difference in arrival date between males and females, with competition among males for access to preferred territories (the rank advantage hypothesis) or mating success (the mate opportunity hypothesis) supposedly driving the evolution of protandry. The fitness costs and benefits of protandry accruing to individuals differing in degree of protandry (arrival date of a male relative to the arrival date of his partner) have never been quantified. We analyzed the fitness consequences of sex differences in arrival date in the barn swallow Hirundo rustica, in which arrival date can be precisely estimated and the fitness of pairs differing in degree of individual protandry assessed. Early arriving males had greater mating success than late arriving males. The number of extrapair offspring in own nests decreased with increasing degree of individual protandry, whereas the number of offspring fathered by a focal male was unrelated to individual protandry. There was directional selection on individual protandry as shown by pairs with a larger than average degree of protandry reproducing early and, hence, supposedly producing more recruits. There was also stabilizing selection on individual protandry as shown by pairs with an intermediate degree of protandry reproducing early. Annual production of fledglings increased with early arrival of males, but not with early arrival of females, once the effect of laying date had been considered, with no additional effect of individual protandry. Neither male nor female survival was significantly related to degree of individual protandry. These findings are consistent not only with the mate opportunity hypothesis but also with a sexual conflict hypothesis, suggesting that males and females differ in their optimal timing of arrival due to sex-specific fitness costs and benefits.