Social groups of gorillas were observed in three captive facilities and one African field site. Cases of potential gesture use, totalling 9,540, were filtered by strict criteria for intentionality, giving a corpus of 5,250 instances of intentional gesture use. This indicated a repertoire of 102 gesture types. Most repertoire differences between individuals and sites were explicable as a consequence of environmental affordances and sampling effects: overall gesture frequency was a good predictor of universality of occurrence. Only one gesture was idiosyncratic to a single individual, and was given only to humans. Indications of cultural learning were few, though not absent. Six gestures appeared to be traditions within single social groups, but overall concordance in repertoires was almost as high between as within social groups. No support was found for the ontogenetic ritualization hypothesis as the chief means of acquisition of gestures. Many gestures whose form ruled out such an origin, i.e. gestures derived from species-typical displays, were used as intentionally and almost as flexibly as gestures whose form was consistent with learning by ritualization. When using both classes of gesture, gorillas paid specific attention to the attentional state of their audience. Thus, it would be unwarranted to divide ape gestural repertoires into 'innate, species-typical, inflexible reactions' and 'individually learned, intentional, flexible communication'. We conclude that gorilla gestural communication is based on a speciestypical repertoire, like those of most other mammalian species but very much larger. Gorilla gestures are not, however, inflexible signals but are employed for intentional communication to specific individuals.

Great apes give gestures deliberately and voluntarily, in order to influence particular target audiences, whose direction of attention they take into account when choosing which type of gesture to use. These facts make the study of ape gesture directly relevant to understanding the evolutionary precursors of human language; here we present an assessment of ape gesture from that perspective, focusing on the work of the “St Andrews Group” of researchers. Intended meanings of ape gestures are relatively few and simple. As with human words, ape gestures often have several distinct meanings, which are effectively disambiguated by behavioural context. Compared to the signalling of most other animals, great ape gestural repertoires are large. Because of this, and the relatively small number of intended meanings they achieve, ape gestures are redundant, with extensive overlaps in meaning. The great majority of gestures are innate, in the sense that the species’ biological inheritance includes the potential to develop each gestural form and use it for a specific range of purposes. Moreover, the phylogenetic origin of many gestures is relatively old, since gestures are extensively shared between different genera in the great ape family. Acquisition of an adult repertoire is a process of first exploring the innate species potential for many gestures and then gradual restriction to a final (active) repertoire that is much smaller. No evidence of syntactic structure has yet been detected.

Great apes give gestures deliberately and voluntarily, in order to influence particular target audiences, whose direction of attention they take into account when choosing which type of gesture to use. These facts make the study of ape gesture directly relevant to understanding the evolutionary precursors of human language; here we present an assessment of ape gesture from that perspective, focusing on the work of the ''St Andrews Group'' of researchers. Intended meanings of ape gestures are relatively few and simple. As with human words, ape gestures often have several distinct meanings, which are effectively disambiguated by behavioural context. Compared to the signalling of most other animals, great ape gestural repertoires are large. Because of this, and the relatively small number of intended meanings they achieve, ape gestures are redundant, with extensive overlaps in meaning. The great majority of gestures are innate, in the sense that the species' biological inheritance includes the potential to develop each gestural form and use it for a specific range of purposes. Moreover, the phylogenetic origin of many gestures is relatively old, since gestures are extensively shared between different genera in the great ape family. Acquisition of an adult repertoire is a process of first exploring the innate species potential for many gestures and then gradual restriction to a final (active) repertoire that is much smaller. No evidence of syntactic structure has yet been detected.

‘Contest hoots’ are acoustically complex vocalisations produced by adult and subadult male bonobos (Pan paniscus). These calls are often directed at specific individuals and regularly combined with gestures and other body signals. The aim of our study was to describe the multi-modal use of this call type and to clarify its communicative and social function. To this end, we observed two large groups of bonobos, which generated a sample of 585 communicative interactions initiated by 10 different males. We found that contest hooting, with or without other associated signals, was produced to challenge and provoke a social reaction in the targeted individual, usually agonistic chase. Interestingly, ‘contest hoots’ were sometimes also used during friendly play. In both contexts, males were highly selective in whom they targeted by preferentially choosing individuals of equal or higher social rank, suggesting that the calls functioned to assert social status. Multi-modal sequences were not more successful in eliciting reactions than contest hoots given alone, but we found a significant difference in the choice of associated gestures between playful and agonistic contexts. During friendly play, contest hoots were significantly more often combined with soft than rough gestures compared to agonistic challenges, while the calls' acoustic structure remained the same. We conclude that contest hoots indicate the signaller's intention to interact socially with important group members, while the gestures provide additional cues concerning the nature of the desired interaction.

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Great ape gestures have attracted considerable research interest in recent years, prompted by their flexible and intentional pattern of use; but almost all studies have focused on single gestures. Here, we report the first quantitative analysis of sequential gesture use in western gorillas (Gorilla gorilla gorilla), using data from three captive groups and one African study site. We found no evidence that gesture sequences were given for reasons of increased communicative efficiency over single gestures. Longer sequences of repeated gestures did not increase the likelihood of response, and using a sequence was seldom in reaction to communicative failure. Sequential combination of two gestures with similar meanings did not generally increase effectiveness, and sometimes reduced it. Gesture sequences were closely associated with play contexts. Markov transition analysis showed two networks of frequently co-occurring gestures, both consisting of gestures used to regulate play. One network comprised only tactile gestures, the other a mix of silent, audible and tactile gestures; apparently, these clusters resulted from gesture use in play with proximal or distal contact, respectively. No evidence was found for syntactic effects of sequential combination: meanings changed little or not at all. Semantically, many gestures overlapped massively with others in their core information (i.e. message), and gesture messages spanned relatively few functions. We suggest that the underlying semantics of gorilla gestures is highly simplified compared to that of human words. Gesture sequences allow continual adjustment of the tempo and nature of social interactions, rather than generally conveying semantically referential information or syntactic structures.

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Summary Many social animals interact jointly, but only humans experience a specific sense of obligation toward their co-participants, a joint commitment . However, joint commitment is not only a mental state but also a process that reveals itself in the coordination efforts deployed during entry and exit phases of joint action. Here, we investigated the presence and duration of such phases in N = 1,242 natural play and grooming interactions of captive chimpanzees and bonobos. The apes frequently exchanged mutual gaze and communicative signals prior to and after engaging in joint activities with conspecifics, demonstrating entry and exit phases comparable to those of human joint activities. Although rank effects were less clear, phases in bonobos were more moderated by friendship compared to phases in chimpanzees, suggesting bonobos were more likely to reflect patterns analogous to human “face management”. This suggests that joint commitment as process was already present in our last common ancestor with Pan .