The underlying causes of differential migration: assumptions, hypotheses, and predictions

Mechanisms governing the migratory decisions of birds have long fascinated ecologists and sparked considerable debate. Identifying factors responsible for variation in migration distance, also known as differential migration, has been a popular approach to understanding the mechanisms underlying migratory behaviour more generally. However, research progress has been slowed by the continued testing of overlapping, non-mechanistic, and circular predictions among a small set of historically entrenched hypotheses. We highlight the body size hypothesis and suggest that the predictions commonly tested have impeded progress because body size relationships with migration distance are predictions made by several distinct hypotheses with contrasting mechanisms. The cost of migration itself has not been adequately accounted for in most hypotheses, and we propose two flight efficiency hypotheses with time- and energy-minimizing mechanisms that allow individuals to mitigate the risks inherent to longer migrations. We also advance two conceptual versions of the social dominance hypothesis based on two distinct underlying mechanisms related to distance minimization and food maximization that will help clarify the role of competition in driving migratory decisions. Overall, we describe and refine 12 mechanistic hypotheses proposed to explain differential migration (along with several other special-case hypotheses), seven of which have underlying mechanisms related to food limitation as past research has identified this to be an important driver of differential migration. We also thoroughly reviewed 145 publications to assess the amount of support for 10 critical assumptions underlying alternative hypotheses for differential migration in birds. Our review reveals that surprisingly few studies explicitly evaluate assumptions within a differential migration context. Generating and testing strong predictions and critical assumptions underlying mechanisms of alternative hypotheses will improve our ability to differentiate among these explanations of differential migration. Additionally, future intraspecific progress will be greatest if investigators continue to focus on mechanisms underlying variation in migration distance within rather than among demographic classes, as previous research has found differing mechanisms to be responsible for differential migration among demographic classes. Interspecifically, a thorough comparative analysis that seeks to explain variation in migration distance among species would broaden both our understanding of the mechanisms regulating current differential migration patterns and those that led to the evolution of migration more generally. Collectively, we provide a framework that, together with advances in animal-borne tracking and other technology, can be used to advance our understanding of the causes of differential migration distance, and migratory decisions more generally.