Size matters but hunger prevails-begging and provisioning rules in blue tit families

It is commonly observed in many bird species that dependent offspring vigorously solicit for food transfers provided by their parents. However, the likelihood of receiving food does not only depend on the parental response, but also on the degree of sibling competition, at least in species where parents raise several offspring simultaneously. To date, little is known about whether and how individual offspring adjusts its begging strategy according to the entwined effects of need, state and competitive ability of itself and its siblings. We here manipulated the hunger levels of either the two heaviest or the two lightest blue tit (Cyanistes caeruleus) nestlings in a short-term food deprivation experiment. Our results showed that the lightest nestlings consistently begged more than the heaviest nestlings, an effect that was overruled by the tremendous increase in begging behaviour after food deprivation. Meanwhile, the amplified begging signals after food deprivation were the only cue for providing parents in their decision process. Furthermore, we observed flexible but state-independent begging behaviour in response to changes in sibling need. As opposed to our expectations, nestlings consistently increased their begging behaviour when confronted with food deprived siblings. Overall, our study highlights that individual begging primarily aims at increasing direct benefits, but nevertheless reflects the complexity of a young birds' family life, in addition to aspects of intrinsic need and state.