Playing with the face: playful facial "chattering" and signal modulation in a monkey species (Theropithecus gelada).

Darwin (1872), in The expression of emotions in man and animals, underlined that human facial expressions represent a shared heritage of our species with nonhuman primates. Play is a fertile field to examine the role of facial expressions that we share with our common ancestors because the primate play face is homologous to human laughter. Here, we focus on the use of two playful expression variants (PF: play face, mouth opened with only the lower teeth exposed; FPF: full play face, lower/upper teeth and gums exposed via the actively retraction of the upper lip) in Theropithecus gelada. During ontogeny PF was replaced by FPF; in older subjects PF was virtually absent. The ontogenetic transition appears to reflect the phylogenetic sequence of the two playful displays with FPF considered a derived form of PF. This age-trend bias of facial displays is probably due to their different roles in communication. The correspondence between facial signals emitted and elicited is a valuable criterion to evaluate playmates' attentional state. Adults were more sensitive than immatures in responding to the play faces of others. Probably, previous playful experience, social competence, and neural circuit maturation are at the basis of adult sensitiveness. Similar to humans, where unconscious laughing is deserved for close friends and/or relatives, FPF was extremely frequent during gelada mother-offspring play. Probably, under some intimate circumstances, facial displays should be primarily linked to the spontaneous expression of emotional states of the sender more than to the strategic transfer of actual information to the receiver.