The function of primate multimodal communication.

Language is commonly narrowed down to speech, but human face-to-face communication is in fact an intrinsically multimodal phenomenon. Despite growing evidence that the communication of non-human primates, our main model for the evolution of language, is also inherently multimodal, most studies on primate communication have focused on either gestures or vocalizations in isolation. Accordingly, the biological function of multimodal signalling remains poorly understood. In this paper, we aim to merge the perspectives of comparative psychology and behavioural ecology on multimodal communication, and review existing studies in great apes for evidence of multimodal signal function based on content-based, efficacy-based and inter-signal interaction hypotheses. We suggest that cross-species comparisons of great ape interactions in both captive and wild settings will allow us to test the conditions in which these hypotheses apply. We expect such studies to provide novel insights into the function of speech-accompanying signals and cues, such as gestures, facial expressions, and eye gaze.