The social role of song in wild zebra finches

Male songbirds sing to establish territories and to attract mates.^1,2 However, increasing reports of singing in non-reproductive contexts³ and by females^4,5 show that song use is more diverse than previously considered. Therefore, alternative functions of song, such as social cohesion³ and synchronization of breeding, by and large, were overlooked even in such well-studied species such as the zebra finch (Taeniopygia guttata). In these social songbirds, only the males sing, and pairs breed synchronously in loose colonies,^6,7 following aseasonal rain events in their arid habitat.^8,9 As males are not territorial, and pairs form long-term monogamous bonds early in life, conventional theory predicts that zebra finches should not sing much at all; however, they do and their song is the focus of hundreds of lab-based studies.^{10,11,12,13,14,15,16,17,18,19,20,21,22} We hypothesize that zebra finch song functions to maintain social cohesion and to synchronize breeding. Here, we test this idea using data from 5 years of field studies, including observational transects, focal and year-round audio recordings, and a large-scale playback experiment. We show that zebra finches frequently sing while in groups, that breeding status influences song output at the nest and at aggregations, that they sing year round, and that they predominantly sing when with their partner, suggesting that the song remains important after pair formation. Our playback reveals that song actively features in social aggregations as it attracts conspecifics. Together, these results demonstrate that birdsong has important functions beyond territoriality and mate choice, illustrating its importance in coordination and cohesion of social units within larger societies.