The songbird as a percussionist: syntactic rules for non-vocal sound and song production in Java sparrows.

Music and dance are two remarkable human characteristics that are closely related. Communication through integrated vocal and motional signals is also common in the courtship displays of birds. The contribution of songbird studies to our understanding of vocal learning has already shed some light on the cognitive underpinnings of musical ability. Moreover, recent pioneering research has begun to show how animals can synchronize their behaviors with external stimuli, like metronome beats. However, few studies have applied such perspectives to unraveling how animals can integrate multimodal communicative signals that have natural functions. Additionally, studies have rarely asked how well these behaviors are learned. With this in mind, here we cast a spotlight on an unusual animal behavior: non-vocal sound production associated with singing in the Java sparrow (Lonchura oryzivora), a songbird. We show that male Java sparrows coordinate their bill-click sounds with the syntax of their song-note sequences, similar to percussionists. Analysis showed that they produced clicks frequently toward the beginning of songs and before/after specific song notes. We also show that bill-clicking patterns are similar between social fathers and their sons, suggesting that these behaviors might be learned from models or linked to learning-based vocalizations. Individuals untutored by conspecifics also exhibited stereotypical bill-clicking patterns in relation to song-note sequence, indicating that while the production of bill clicking itself is intrinsic, its syncopation appears to develop with songs. This paints an intriguing picture in which non-vocal sounds are integrated with vocal courtship signals in a songbird, a model that we expect will contribute to the further understanding of multimodal communication.