Sequential settlement and site dependence in a migratory raptor

Habitat selection models, such as the ideal free, ideal despotic, site-dependent, and conspecific cuing models, are of great importance to behavioral ecologists given their capability to predict habitat distributions and to link individual behavior to population processes. However, there have been relatively few field tests of their predictions. We tested the 4 models by studying the process of sequential settlement on territory in 2 distant populations of a migratory raptor, the black kite Milvus migrans. Results were mainly consistent with the site-dependent model: on arrival, kites settled on progressively lower quality territories, and earlier arriving individuals were older, larger, and in better body condition than later arriving ones, leading to a statedependent arrival sequence also predicted by a previous theoretical model of settlement pattern. Occupation of superior territories by superior phenotypes resulted in cascading advantages for earlier arriving individuals in terms of subsequent reproductive performance. At the population level, the populations expanded/retracted from lower quality sites during population increases/declines. The above scenario was consistent across the 2 populations, and a review of the literature uncovered a remarkably consistent picture of state-dependent arrival, progressive monopolization of best quality sites, and cascading effects on subsequent breeding performance. We propose as a general paradigm of sequential settlement the following process: 1) arrival date is a reliable surrogate of phenotype quality, 2) early-arriving individuals have preferential access to the best quality sites and partners, 3) the above conditions cascade into a number of benefits ultimately related to higher fitness for earlier arriving individuals.