Foraging intention affects whether willow tits call to attract members of mixed-species flocks

Understanding how individual behaviour influences the spatial and temporal distribution of other species is necessary to resolve the complex structure of species assemblages. Mixed-species bird flocks provide an ideal opportunity to investigate this issue, because members of the flocks are involved in a variety of behavioural interactions between species. Willow tits (Poecile montanus) often produce loud calls when visiting a new foraging patch to recruit other members of mixed-species flocks. The costs and benefits of flocking would differ with individual foraging behaviours (i.e. immediate consumption or caching); thus, willow tits may adjust the production of loud calls according to their foraging intention. In this study, we investigated the link between foraging decisions and calling behaviour in willow tits and tested its influence on the temporal cohesion with members of mixed-species flocks. Observations at experimental foraging patches showed that willow tits produced more calls when they consumed food items compared with when they cached them. Playback experiments revealed that these calls attracted flock members and helped to maintain their presence at foraging patches. Thus, willow tits adjusted calling behaviour according to their foraging intention, thereby coordinating the associations with members of mixed-species flocks. Our findings demonstrate the influence of individual decision-making on temporal cohesion with other species and highlight the importance of interspecific communication in mixed-species flocking dynamics.