Adaptive hypotheses for protandry in arrival to breeding areas: a review of models and empirical tests

Abstract

Our understanding of avian migration has progressed significantly, yet the selective conditions that favor the arrival of males before females at the site of reproduction remain largely unclear. Here, we review the leading adaptive hypotheses for protandry, highlight some key empirical studies that test protandry theory, and identify theoretical and empirical information demands. In general, protandry should evolve in species where the variance in male reproductive success is larger than in females if the costs to males of earlier arrival relative to calendar date (viability selection) can be balanced by increased mating opportunities (sexual selection). Early arrival by males can provide ‘priority benefits’ that help in the monopolization of resources or ‘early bird draw benefits’ that increase opportunities for extra-pair mating. While some empirical studies are consistent with theoretical predictions regarding the important selection factors that influence protandry (e.g., extrinsic mortality and extra-pair paternity), some are not, and some studies focus on ecological factors that have not been considered explicitly by theory. We call for an integrated theoretical approach to help formalize how protandry should evolve in response to the antagonistic roles of natural and sexual selection, the nature of competitive asymmetries among males or females, sex-specific costs and benefits of early arrival, and various climate change scenarios.