Predictors of floater status in a long-lived bird: a cross-sectional and longitudinal test of hypotheses

Few studies have been capable of monitoring the nonterritorial sector of a population because of the typically secretive behaviour of floating individuals, despite the existing consensus over the demographic importance of floating. Furthermore, there is almost no information on floating behaviour for migratory species. The factors that determine whether an individual will be a floater or a territory owner have been framed into five, non-mutually exclusive hypotheses: (i) territory holders are morphologically superior to floaters (resource-holding potential hypothesis); (ii) age confers skills and fighting motivation which lead to social dominance and territory ownership (age hypothesis); (iii) occupancy time of a site determines asymmetries in its knowledge, familiarity and value for potential contenders (site-dominance hypothesis); (iv) contenders use an arbitrary rule to settle contests leading to pre-defined cut-off points for a biologically meaningful trait (e.g. age, body size) separating floaters from territory holders (arbitrary convention hypothesis); and (v) floaters set up a 'war of attrition' at arbitrarily chosen territories (arbitrary attrition hypothesis). We tested these hypotheses using long-term data on a long-lived, migratory raptor, the black kite Milvus migrans Boddaert. Floating status was best explained by the concerted action of mechanisms consistent with the age and site-dominance hypotheses. In both cross-sectional and longitudinal analyses, acquisition of a territory was determined by a complex interaction between age and early arrival from migration, suggesting: (i) a progressive incorporation of early arriving individuals in the territorial contingent of the population, and (ii) the existence of an alternative restraint strategy of delayed territoriality mediated by long-term acquisition of social dominance. Such results suggested that territory acquisition was mediated by the establishment of site dominance through pre-emption and, secondarily, despotism. In this population, age and arrival date aligned individuals along a demographic continuum ranging from successful breeders monopolizing high-quality resources to floaters with no resources, consistent with the notion of floating as an extreme form of breeding failure.