Vocal-visual combinations in wild chimpanzees

Human communication is strikingly multi-modal, relying on vocal utterances combined with visual gestures, facial expressions and more. Recent efforts to describe multi-modal signal production in our ape relatives have shed important light on the evolutionary trajectory of this core hallmark of human language. However, whilst promising, a systematic quantification of primate signal production which filters out random combinations produced across modalities is currently lacking. Here, through recording the communicative behaviour of wild chimpanzees from the Kibale forest, Uganda we address this issue and generate the first repertoire of non-random combined vocal and visual components. Using collocation analysis, we identify more than 100 vocal-visual combinations which occur more frequently than expected by chance. We also probe how multi-modal production varies in the population, finding no differences between individuals as a function of age, sex or rank. The number of visual components exhibited alongside vocalizations was, however, associated with vocalization type and duration. We demonstrate that chimpanzees produce a vast array of combined vocal and visual components, exhibiting a hitherto underappreciated level of combinatorial complexity. We conclude that a multi-modal approach is crucial to accurately representing the communicative abilities of non-human primates.