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Rapid facial mimicry (RFM), the rapid and automatic replication of facial expression perceived, is considered a basic form of empathy and was investigated mainly during play. RFM occurs in Catarrhini (Old World primates), but it is not still demonstrated in Platyrrhini (New World primates). For this reason, we collected video data on playful interactions (Nplay_interactions = 149) in three species of spider monkeys (Ateles fusciceps-N = 11, Ateles hybridus-N = 14, and Ateles paniscus-N = 6) housed at La Vallée des Singes and the ZooParc de Beauval (France). For the first time, we demonstrated the occurrence of RFM in Platyrrhini (analyzing 175 events). Players' sex, age, species, relationship quality, and kinship did not modulate RFM probably due to the species' complex fission-fusion dynamics and flexible interindividual social relationships. Compared to the absence of any playful expressions or the presence of only not replicated play face, RFM prolonged the session duration and was sequentially associated with more types of more intense offensive playful patterns (patterns aimed at attacking/pursuing the playmate). We proposed that RFM may favor synchronization and context sharing between players, thus decreasing the risk of behavior misinterpretation while simultaneously fostering a more competitive nature of play. In conclusion, this study stimulates additional research on the evolutionary origins of motor mimicry in primates, possibly dating back to before the divergence of New and Old World monkeys. Furthermore, it also points toward the possibility that RFM may not always lead to cooperation but also to competition, depending on the context and species' social and cognitive features.

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Animals often initiate social interactions by exchanging signals. Especially when initiating amicable interactions, signaling one's friendly stance toward others in advance may be important to avoid being misunderstood as having hostile intentions. We used data on dyadic play fighting in a group of Japanese macaques, Macaca fuscata, to examine the function of "play face" at the opening of a play session. We found no support for the previously proposed hypothesis that play face expression is likelier before entering risky situations (e.g., before gaining an undue advantage over the partner) to avoid being misunderstood. The results showed that play face expression was likelier in male juveniles before initiating play with other males than in females before initiating play with males or other females and that juveniles were likelier to express play face before initiating play with others closer in age. As male Japanese macaques play more frequently than females, and juveniles prefer to play with individuals closer in age, play face expression before play initiation may reflect the individual's motivation for subsequent play interactions. This interpretation is supported by our observation that play bouts lasted longer when initiated with bidirectional play face by both participants than when initiated without play face. We also argued that since there was no tendency that play face was likelier to be expressed toward individuals with low play propensity (e.g., females) or infrequent partners to play with (e.g., individuals more distant in age), Japanese macaques may not tactically deploy this signal to recruit reluctant partners.

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Among African great apes, play is virtually absent between adult lowland gorillas (Gorilla gorilla gorilla). Here, we report an extremely rare case of adult-adult play observed in the lowland gorilla group housed at La Vallée de Singes (France). We recorded three playful interactions between the silverback and an adult lactating female. Given the diverse causal and functional nature of play, different factors may join in promoting this behaviour. In our group, contrary to what has been shown by previous studies in wild and captive gorillas, adult females spent more time in spatial proximity with the silverback than with other females. Hence, the probability of social interaction (including play) between silverback and adult females was enhanced. Moreover, the motivation of the lactating female to play might be an effect of oxytocin, a hormone that reaches high concentration levels during lactation and that promotes social affiliation. The fact that play invitations were always performed by the female might support this hypothesis. Certainly, we cannot exclude the possibility that the play event is a group idiosyncrasy or an artefact of captivity, even though the subjects never showed abnormal behaviour. Structurally, play sessions showed a suitable degree of pattern variability and switching frequency from one pattern to another. The proportion of offensive patterns was higher in the female during play and in the male during aggression, which conforms to the role reversal play phenomenon. In conclusion, this report confirms that the absence of evidence is not the evidence of absence. It is likely that under particular physiological or socio-ecological conditions, adult-adult play may be manifested as an "unconventional" part of gorilla social behaviour.

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Play is widespread across mammalian taxa, but species strongly vary in the ways they play. In less despotic primate species (i.e., with less steep dominance hierarchies, less severe conflicts, and more reconciliation), play has been described as being more frequent, cooperative, and freely expressed. To study the link between social play and dominance style, we compared play behavior in free-ranging infants, juveniles and subadults of more despotic Japanese macaques (Macaca fuscata, N = 24) and less despotic moor macaques (Macaca maura, N = 17). We found interspecific differences in play behavior that corresponded with the contrasting dominance styles of the study species, largely confirming our predictions. In particular, moor macaques spent a larger proportion of time in solitary and social play than Japanese macaques, while Japanese macaques spent a larger proportion of time in grooming interactions. In moor macaques, play sessions included more players, a larger variety of play behaviors, greater play face rates, a greater proportion of time in contact play, and a higher rate of reciprocal play-biting than in Japanese macaques. Aggressive escalations were not common, but more frequent in Japanese macaques. Finally, a higher frequency of play faces during play sessions predicted the occurrence of more reciprocal play-bites, but not the proportion of time spent in contact play behaviors. Additional studies on other groups and species will allow a better understanding of the link between dominance style and social play.

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Spontaneous mimicry appears fundamental to emotional perception and contagion, especially when it involves facial emotional expressions. Here we cover recent evidence on spontaneous mimicry from ethology, psychology and neuroscience, in non-human and human animals. We first consider how mimicry unfolds in non-human animals (particularly primates) and how it relates to emotional contagion. We focus on two forms of mimicry-related phenomena: facial mimicry and yawn contagion, which are largely conserved across mammals and useful to draw evolutionary scenarios. Next, we expand on the psychological evidence from humans that bears on current theoretical debates and also informs non-human animal research. Finally, we cover the neural bases of facial mimicry and yawn contagion. We move beyond the perception/expression/experience trichotomy and from the correlational to the causal evidence that links facial mimicry to emotional contagion by presenting evidence from neuroimaging, direct manipulation, neuro-stimulation and lesion studies. In conclusion, this review proposes a bottom-up, multidisciplinary approach to the study of spontaneous mimicry that accounts for the evolutionary continuity linking non-human and human animals.

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Play behaviors and signals during playful interactions with juvenile conspecifics are important for both the social and cognitive development of young animals. The social organization of a species can also influence juvenile social play. We examined the relationships among play behaviors, candidate play signals, and play bout termination in Tibetan macaques (Macaca thibetana) during juvenile and infant social play to characterize the species play style. As Tibetan macaques are despotic and live in groups with strict linear dominance hierarchies and infrequent reconciliation, we predicted that play would be at risk of misinterpretation by both the individuals engaged in the play bout and by those watching, possibly leading to injury of the players. Animals living in such societies might need to frequently and clearly signal playful intent to play partners and other group members to avoid aggressive outcomes. We gathered video data on 21 individually-identified juvenile and infant macaques (one month to five years of age) from the Valley of the Wild Monkeys, Mt. Huangshan, China. We used all-occurrence sampling to record play behaviors and candidate play signals based on an ethogram. We predicted that play groups would use multiple candidate play signals in a variety of contexts and in association with the number of audience members in proximity to the players and play bout length. In the 283 playful interactions we scored, juvenile and infant macaques used multiple body and facial candidate play signals. Our data showed that juvenile and infant Tibetan macaques use a versatile repertoire of play behaviors and signals to sustain play.

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Knowledge of the context and development of playful expressions in chimpanzees is limited because research has tended to focus on social play, on older subjects, and on the communicative signaling function of expressions. Here we explore the rate of playful facial and body expressions in solitary and social play, changes from 12- to 15-months of age, and the extent to which social partners match expressions, which may illuminate a route through which context influences expression. Naturalistic observations of seven chimpanzee infants (Pan troglodytes) were conducted at Chester Zoo, UK (n = 4), and Primate Research Institute, Japan (n = 3), and at two ages, 12 months and 15 months. No group or age differences were found in the rate of infant playful expressions. However, modalities of playful expression varied with type of play: in social play, the rate of play faces was high, whereas in solitary play, the rate of body expressions was high. Among the most frequent types of play, mild contact social play had the highest rates of play faces and multi-modal expressions (often play faces with hitting). Social partners matched both infant play faces and infant body expressions, but play faces were matched at a significantly higher rate that increased with age. Matched expression rates were highest when playing with peers despite infant expressiveness being highest when playing with older chimpanzees. Given that playful expressions emerge early in life and continue to occur in solitary contexts through the second year of life, we suggest that the play face and certain body behaviors are emotional expressions of joy, and that such expressions develop additional social functions through interactions with peers and older social partners.