Windscape and tortuosity shape the flight costs of northern gannets.

When animals move across a landscape, they alternate between active searching phases in areas with high prey density and commuting phases towards and in-between profitable feeding patches. Such active searching movements are more sinuous than travelling movements, and supposedly more costly in energy. Here we provide an empirical validation of this long-lasting assumption. To this end, we evaluated simultaneously energy expenditure and trajectory in northern gannets (Morus bassanus) using GPS loggers, dive recorders and three-dimensional accelerometers. Three behavioural states were determined from GPS data: foraging, when birds actively searched for prey (high tortuosity, medium speed); travelling, when birds were commuting (straight trajectory, high speed); and resting (straight trajectory, low speed). Overall dynamic body acceleration, calculated from acceleration data, was used as a proxy for energy expenditure during flight. The impact of windscape characteristics (wind force and direction) upon flight costs was also tested. Energy expenditure of northern gannets was higher during sinuous foraging flight than during more rectilinear travelling flight, demonstrating that turns are indeed costly. Yet wind force and direction also strongly shaped flight energy expenditure; within any behavioural state it was less costly to fly with the wind than against it, and less costly to fly with strong winds. Despite the major flight costs of wind action, birds did not fully optimize their flight track relative to wind direction, probably because of prey distributions relative to the coastline and wind predictability. Our study illustrates how both tortuosity and windscape shape the foraging costs of marine predators such as northern gannets.