Factors influencing route choice by avian migrants: a dynamic programming model of Pacific brant migration

We used stochastic dynamic programming to investigate a spectacular migration strategy in the black brant Branta bernicla nigricans, a species of goose. Black brant migration is well suited for theoretical analysis since there are a number of existing strategies that easily can be compared. In early autumn, almost the entire population of the black brant gathers at Izembek Lagoon on the Alaska Peninsula to stage and refuel before the southward migration. There are at least three distinct strategies, with most geese making a spectacular direct migration more than 5000km across the Gulf of Alaska to their wintering grounds in southern Baja California or mainland Mexico. This is a potentially dangerous strategy since foraging is not possible during the overseas passage. Some individuals instead use shorter flights to make a detour along the coast, a longer route that all individuals use for northwards migration in spring. Since flight costs accelerate with increasing body mass, migration by short flights is energetically cheaper than long-distance flights. A small but increasing part of the population has recently begun to winter at Izembek. We investigated this migration under two different suppositions using a dynamic state variable model. First, if the geese are free to make a strategic choice, under what assumptions should they prefer direct migration and under what assumptions should they prefer detour migration/winter residency? Second, provided that the dominating direct migration strategy is optimal, what conditions will force the geese to go for detour migration/winter residency? In the second case the geese may try to follow an optimal direct migration strategy, but stochastic events may force them to choose a suboptimal policy. We also simulated possible effects of global warming. The model suggests that the fuel level at arrival in Izembek and fuel gain rates are key factors and that tail winds must have been reliable in the past, otherwise direct migration could not have evolved. It also suggests that a change to milder winters may promote an unexpectedly abrupt change from long-distance to short-distance migration or winter residency. Finally, it produced a number of predictions that might be testable in the field.