Multimodal communication and audience directedness in the greeting behaviour of semi-captive African savannah elephants

Many species communicate by combining signals into multimodal combinations. Elephants live in multi-level societies where individuals regularly separate and reunite. Upon reunion, elephants often engage in elaborate greeting rituals where they use vocalisations and body acts produced with different articulators and of various sensory modalities. However, whether elephants use audience directed gestures and how elephants combine vocalisations and gestures during greeting is still unknown. Here we use separation-reunion events to explore the greeting behaviour of semi-captive elephants (Loxodonta africana). We apply Linear Mixed Models and Collocation Analyses to investigate whether elephants direct silent-visual, audible, and tactile gestures at their audience based on their attentional state and how they combine these gestures with vocalisations in multicomponent combinations. We show that elephants select gesture modality according to the audience´s visual attention, indicating evidence of first-order intentional use. We further show that elephants incorporate vocalisations and gestures in different combinations and orders. The most frequent combination consists of rumble vocalisations with ear-flapping gestures, used most often between females. By showing that a species evolutionarily distant to our own primate lineage shows sensitivity to their audience’s visual attention in their gesturing and combines gestures with vocalisations, our study advances our understanding of the emergence of first-order intentionality and multicomponent signalling across taxa.