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Food calling in wild ravens (
 Corvus corax
 ) revisited: Who is addressed?

Numerous birds and mammals use vocal signals to advertise feeding opportunities but often such signals vary with individual and contextual factors. Non-breeding ravens call at food that is difficult to access, resulting in the attraction of nearby conspecifics. Although callers may benefit from group formation in various ways, we recently found substantial individual variation in food calling. We here explored whether this variation can be partly explained by the social dynamics in raven foraging groups, together with already known effects of age class and sex. Specifically, we expected ravens to respond to the presence or absence of affiliates that could act as cooperative partners in the forthcoming feeding event, that is they should call when other ravens were present but they themselves were alone rather than when they were also in company of an affiliation partner. We observed the vocal behaviour of individually marked wild ravens and, simultaneously, categorized their affiliative behaviour with other ravens in the minutes before experimentally controlled feedings. In line with our prediction, individuals were less likely to produce food-associated calls when they were in close contact with an affiliation partner prior to feeding as compared to when they were alone. Furthermore, sex and age class influenced food calling as females called more often than males and younger birds called more often than adult ravens. In conclusion, these results suggest that ravens attempt to find support from a particular cooperative partner by broadly advertise feeding opportunities via food-associated calls, especially when they have low chances in contest competition due to their age and sex. These findings lend further support to the assumption of raven flocks being structured by social relationships and individual birds flexibly controlling their vocal signalling according to the current flock composition.