Accounting for central place foraging constraints in habitat selection studies

Habitat selection studies contrast actual space use with the expected use under the null hypothesis of no selection (hereafter neutral use). Neutral use is most often equated to the relative frequencies with which environmental features occur. This generates a considerable bias when studying habitat selection by foragers that perform numerous trips back and forth to a central place (CP). Indeed, the increased space use close to the CP with respect to distant places reflects a mechanical effect, rather than a true selection for the closest habitats. Yet, correctly estimating habitat selection by CP foragers is of paramount importance for a better understanding of their ecology and to properly plan conservation actions. We show that including the distance to the CP as a covariate in unconditional Resource Selection Functions, as applied in several studies, is ineffective to correct for the bias. This bias can be eliminated only by contrasting the actual use to an appropriate neutral use that considers the CP forager behavior. We also show that the need to specify an appropriate neutral use overall distribution can be bypassed by relying on a conditional approach, where the neutral use is assessed locally regardless of the distance to the CP.