Helping as a signal: does removal of potential audiences alter helper behavior in the bell miner?

Despite many studies on the evolution of cooperative breeding and helping at the nest, relatively few have explored the possibility that helping functions as a signal to gain social advantage within groups (the pay to stay and social prestige hypotheses). One of the most promising candidates for such a signaling system is the cooperatively breeding bell miner, Manorina melanophrys. Large numbers of unrelated helpers attend multiple nests while giving individually identifiable vocalizations, breeding females usually remain within monitoring distance of the nest area, and females often re-pair with the hardest working male helper after the death/removal of their breeding partner. We examined the possibility that helping operates as a signal by temporarily removing the potential audience: the breeding male or the breeding female. However, there was no discernable change in provisioning behavior of helpers, relative to control periods. We also simulated the presence of the removed birds through playbacks of their individual-specific calls and again found no effect on others' visit rates, prey types, load size, and a variety of other behaviors at the nest. If either signaling hypothesis explained helping in this system, we might have expected facultative decreases in conspicuous provisioning behaviors when one or other potential audience was absent. Thus, despite possessing many of the prerequisites of a signal-based helping system, there is no evidence for such a phenomenon in bell miners. Cooperation in these groups of mixed relatedness may instead be driven by a combination of kin selection and direct benefits via group augmentation and/or pseudoreciprocity.