Conspecific aggression strategies are conditioned by environmental, social and intrinsic variables in breeding blue tits Cyanistes caeruleus

Territorial behaviour arises as a strategy of ensuring individuals’ access to a variety of potentially limiting resources. While aggressiveness is a well-studied widespread trait across taxa, the mechanisms that allow for a range of aggressive phenotypes to coexist in the wild remains unclear. In this study, we analyse environmental, social and intrinsic variables that can modulate the expression of different strategies of male–male aggressiveness. Furthermore, through network analysis we explore the role of this trait in the establishment of territories during the breeding season as the intensity of different aggressiveness strategies may limit or grant access to resources. Simulating territorial intrusions during the early incubation period, we assessed the aggressiveness of breeding male blue tits (Cyanistes caeruleus). We defined three types of conspecific aggressiveness (nonconfrontational intimidating, nonconfrontational cautious and confrontational) and analysed the effect of habitat structure, territory quality, presence of other breeding species and male condition on the type and intensity of the aggressive display. The results obtained suggest that yearling males rely on intimidating behaviour more than older males, that perform more cautious displays. Furthermore, smaller and heavier males opted for confrontational strategies. The density and nature of neighbours, as well as the territory quality and the habitat structure, also conditioned the intensity and type of display. Surprisingly, the network analysis revealed that the intensity of male–male aggressive displays did not condition the establishment of breeding territories. Our results suggest that aggressiveness is a context-specific trait shaped by a complex array of environmental and intrinsic parameters.