The eyes and ears are visual indicators of attention in domestic horses

Sensitivity to the attentional states of others has adaptive advantages [1] , and in social animals, attending to others is important for predator detection, as well as a pre-requisite for normal social functioning and more complex socio-cognitive abilities [2] . Despite widespread interest in how social species perceive attention in others, studies of non-human animals have been inconclusive about the detailed cues involved [3] . Previous work has focused on head and eye direction, overlooking the fact that many mammals have obvious and mobile ears that could act as a visual cue to attention. Here we report that horses use the head orientation of a conspecific to locate food, but that this ability is disrupted when parts of the face (the eyes and ears) are covered up with naturalistic masks. The ability to correctly judge attention also interacted with the identity of the model horse, suggesting that individual differences in facial features may influence the salience of cues. Our results indicate that a combination of head orientation with facial expression, specifically involving both the eyes and ears, is necessary for communicating social attention. These findings emphasise that in order to understand how attention is communicated in non-human animals, it is essential to consider a broad range of cues