Anna Ilona Roberts

Communicative intentions in wild chimpanzees: persistence and elaboration in gestural signalling

We examine evidence for communicative intent during conspecific interactions in wild chimpanzees (Budongo Forest, Uganda), focusing on persistence in gestural communication. Previous research indicates that great apes have large gestural repertoires and produce gestural communication in a flexible and intentional manner, including the production of gesture sequences. Although there is a lack of consensus on the form and function of sequences, there is some evidence that sequences are produced when signallers fail to receive any response from a recipient. Here, we provide first systematic evidence for communicative persistence in wild chimpanzees. Rather than examining only the presence or absence of a response, we used the most commonly observed response to assign meanings to gestures and examined sequence production in relation to response congruency. Chimpanzees ceased communication if successful, but persevered when unsuccessful. Chimpanzees repeated gestures when a response partially matched their goal but substituted the original gesture when a response was incongruent. Persistence was also mediated by recipient intent to respond, with more sequences produced within competitive than affiliative contexts. Gestures within sequences were homogenous in semantic meaning and signallers continued until the response matched the assigned meaning of the initial gesture. Gestural sequence production was not primarily affective; gesture intensity (in terms of modality) did not increase within sequences. Chimpanzee gestural sequences emerged to achieve specific outcomes; given variability in recipient behaviour following initial gestures, signallers were flexible in their persistence towards these goals.

Usage and comprehension of manual gestures in wild chimpanzees

Flexibility is considered a defining feature of great ape gestural communication. Previous research has suggested that there is a ‘means–ends’ dissociation between gesture type and context, whereby one signal may be used across contexts and several signals used within the same context. Such flexibility in signal production demands contextual comprehension, whereby recipients may perceive the context-free message of a given manual gesture, but also decide how to respond by inferring the signallers goals from the accompanying context. We conducted naturalistic observations of wild East African chimpanzee, Pan troglodytes schweinfurthii, manual gestures, focusing on recipient perspective- during communicative interactions. Our results indicate that chimpanzees recognize the context-free meanings of gestures and they are also able to respond flexibly by inferring the meaning from the combination of gesture and context, including relative rank. When analysed at the level of gesture type, some gestures were tightly associated with dominant responses and outcomes. Chimpanzee manual gestures are primarily used for directing a recipients movement or attention but the motivation underlying these gestural requests is inferred by the recipient from the context.

Chimpanzees modify intentional gestures to coordinate a search for hidden food

Humans routinely communicate to coordinate their activities, persisting and elaborating signals to pursue goals that cannot be accomplished individually. Communicative persistence is associated with complex cognitive skills such as intentionality, because interactants modify their communication in response to anothers understanding of their meaning. Here we show that two language-trained chimpanzees effectively use intentional gestures to coordinate with an experimentally-naïve human to retrieve hidden food, providing some of the most compelling evidence to date for the role of communicative flexibility in successful coordination in nonhumans. Both chimpanzees (named Panzee and Sherman) increase the rate of non-indicative gestures when the experimenter approaches the location of the hidden food. Panzee also elaborates her gestures in relation to the experimenters pointing, which enables her to find food more effectively than Sherman. Communicative persistence facilitates effective communication during behavioural coordination and is likely to have been important in shaping language evolution.

The repertoire and intentionality of gestural communication in wild chimpanzees

A growing body of evidence suggests that human language may have emerged primarily in the gestural rather than vocal domain, and that studying gestural communication in great apes is crucial to understanding language evolution. Although manual and bodily gestures are considered distinct at a neural level, there has been very limited consideration of potential differences at a behavioural level. In this study, we conducted naturalistic observations of adult wild East African chimpanzees (Pan troglodytes schweinfurthii) in order to establish a repertoire of gestures, and examine intentionality of gesture production, use and comprehension, comparing across manual and bodily gestures. At the population level, 120 distinct gesture types were identified, consisting of 65 manual gestures and 55 bodily gestures. Both bodily and manual gestures were used intentionally and effectively to attain specific goals, by signallers who were sensitive to recipient attention. However, manual gestures differed from bodily gestures in terms of communicative persistence, indicating a qualitatively different form of behavioural flexibility in achieving goals. Both repertoire size and frequency of manual gesturing were more affiliative than bodily gestures, while bodily gestures were more antagonistic. These results indicate that manual gestures may have played a significant role in the emergence of increased flexibility in great ape communication and social bonding.

Wild chimpanzees modify modality of gestures according to the strength of social bonds and personal network size

Primates form strong and enduring social bonds with others and these bonds have important fitness consequences. However, how different types of communication are associated with different types of social bonds is poorly understood. Wild chimpanzees have a large repertoire of gestures, from visual gestures to tactile and auditory gestures. We used social network analysis to examine the association between proximity bonds (time spent in close proximity) and rates of gestural communication in pairs of chimpanzees when the intended recipient was within 10 m of the signaller. Pairs of chimpanzees with strong proximity bonds had higher rates of visual gestures, but lower rates of auditory long-range and tactile gestures. However, individual chimpanzees that had a larger number of proximity bonds had higher rates of auditory and tactile gestures and lower rates of visual gestures. These results suggest that visual gestures may be an efficient way to communicate with a small number of regular interaction partners, but that tactile and auditory gestures may be more effective at communicating with larger numbers of weaker bonds. Increasing flexibility of communication may have played an important role in managing differentiated social relationships in groups of increasing size and complexity in both primate and human evolution.

Gestural Communication and Mating Tactics in Wild Chimpanzees.

The extent to which primates can flexibly adjust the production of gestural communication according to the presence and visual attention of the audience provides key insights into the social cognition underpinning gestural communication, such as an understanding of third party relationships. Gestures given in a mating context provide an ideal area for examining this flexibility, as frequently the interests of a male signaller, a female recipient and a rival male bystander conflict. Dominant chimpanzee males seek to monopolize matings, but subordinate males may use gestural communication flexibly to achieve matings despite their low rank. Here we show that the production of mating gestures in wild male East African chimpanzees (Pan troglodytes schweunfurthii) was influenced by a conflict of interest with females, which in turn was influenced by the presence and visual attention of rival males. When the conflict of interest was low (the rival male was present and looking away), chimpanzees used visual/ tactile gestures over auditory gestures. However, when the conflict of interest was high (the rival male was absent, or was present and looking at the signaller) chimpanzees used auditory gestures over visual/ tactile gestures. Further, the production of mating gestures was more common when the number of oestrous and non-oestrus females in the party increased, when the female was visually perceptive and when there was no wind. Females played an active role in mating behaviour, approaching for copulations more often when the number of oestrus females in the party increased and when the rival male was absent, or was present and looking away. Examining how social and ecological factors affect mating tactics in primates may thus contribute to understanding the previously unexplained reproductive success of subordinate male chimpanzees.

Visual Attention, Indicative Gestures, and Calls Accompanying Gestural Communication Are Associated With Sociality in Wild Chimpanzees (Pan troglodytes schweinfurthii)

The challenges of life in complex social groups may select for complex communication to regulate interactions among conspecifics. Whereas the association between social living and vocalizations has been explored in nonhuman primates, great apes also have a rich repertoire of gestures, and how the complexity of gestural communication relates to sociality is still unclear. We used social network analysis to examine the relationship between the duration of time pairs of chimpanzees spent in proximity (within 10 m) and the rates of gestural communication accompanied by visual attention of the signaler, one-to-one calls, indicative gestures (collectively self-relevance cues), and synchronized pant-hoot calls. Pairs of chimpanzees that spent a longer duration of time in proximity had a higher rate of visual gestures accompanied by these behaviors. Further, individual chimpanzees that had a greater number of proximity bonds had a larger social network maintained through gestures accompanied by synchronized pant-hoot calls. In contrast, gestures unaccompanied by these behaviors were not positively associated with either proximity bonds in pairs of chimpanzees or individual differences in sociality. These results suggest that self-relevance cues and synchronized pant-hoot calls accompanying gestures may increase the efficiency of gestural communication in social bonding and that multimodal communication may have played a key role in language evolution.

Convergence and divergence in gesture repertoires as an adaptive mechanism for social bonding in primates

A key challenge for primates living in large, stable social groups is managing social relationships. Chimpanzee gestures may act as a time-efficient social bonding mechanism, and the presence (homogeneity) and absence (heterogeneity) of overlap in repertoires in particular may play an important role in social bonding. However, how homogeneity and heterogeneity in the gestural repertoire of primates relate to social interaction is poorly understood. We used social network analysis and generalized linear mixed modelling to examine this question in wild chimpanzees. The repertoire size of both homogeneous and heterogeneous visual, tactile and auditory gestures was associated with the duration of time spent in social bonding behaviour, centrality in the social bonding network and demography. The audience size of partners who displayed similar or different characteristics to the signaller (e.g. same or opposite age or sex category) also influenced the use of homogeneous and heterogeneous gestures. Homogeneous and heterogeneous gestures were differentially associated with the presence of emotional reactions in response to the gesture and the presence of a change in the recipients behaviour. Homogeneity and heterogeneity of gestural communication play a key role in maintaining a differentiated set of strong and weak social relationships in complex, multilevel societies.

Intentional gestures predict complex sociality in wild chimpanzee

A key challenge for primates is coordinating behavior with conspecifics in large, complex social groups. Gestures play a key role in this process and chimpanzees show considerable flexibility communicating through single gestures, sequences of gestures interspersed with periods of response waiting (persistence) and rapid sequences where gestures are made in quick succession, too rapid for the response waiting to have occurred. Previous studies examined behavioral reactions to single gestures and sequences, but whether this complexity is associated with more complex sociality at the level of the dyad partner and the group as a whole is not well understood. We used social network analysis to examine how the production of single gestures and sequences of gestures was related to the duration of time spent in proximity and individual differences in proximity in wild East African chimpanzees (Pan troglodytes schweinfurthii). Pairs of chimpanzees that spent a longer duration of time in proximity had higher rates of persistence, but not a higher rate of single gesture or rapid sequences. Central individuals in the social network received higher rates of persistence, but not rapid sequence or single gesture. Intentional gestural communication plays an important role in regulating social interactions in complex primate societies.

Persistence in gestural communication predicts sociality in wild chimpanzees

A key challenge for primates is coordinating behaviour with conspecifics in large, complex social groups. Gestures play a key role in this process and chimpanzees show considerable flexibility communicating through single gestures, sequences of gestures interspersed with periods of response waiting (persistence), and rapid sequences where gestures are made in quick succession, too rapid for the response waiting to have occurred. The previous studies examined behavioural reactions to single gestures and sequences, but whether this complexity is associated with more complex sociality at the level of the dyad partner and the group as a whole is not well understood. We used social network analysis to examine how the production of single gestures and sequences of gestures was related to the duration of time spent in proximity and individual differences in proximity in wild East African chimpanzees (Pan troglodytes schweinfurthii). Pairs of chimpanzees that spent a longer duration of time in proximity had higher rates of persistence sequences, but not a higher rate of single gestures or rapid sequences. The duration of time spent in proximity was also related to the rate of responding to gestures, and response to gesture by activity change. These results suggest that communicative persistence and the type of response to gestures may play an important role in regulating social interactions in primate societies.