Abstract
A key driver of brain evolution in primates and humans is the cognitive demands arising from managing social relationships. In primates, grooming plays a key role in maintaining these relationships, but the time that can be devoted to grooming is inherently limited. Communication may act as an additional, more time-efficient bonding mechanism to grooming, but how patterns of communication are related to patterns of sociality is still poorly understood. We used social network analysis to examine the associations between close proximity (duration of time spent within 10 m per hour spent in the same party), grooming, vocal communication, and gestural communication (duration of time and frequency of behavior per hour spent within 10 m) in wild chimpanzees. This study examined hypotheses formulated a priori and the results were not corrected for multiple testing. Chimpanzees had differentiated social relationships, with focal chimpanzees maintaining some level of proximity to almost all group members, but directing gestures at and grooming with a smaller number of preferred social partners. Pairs of chimpanzees that had high levels of close proximity had higher rates of grooming. Importantly, higher rates of gestural communication were also positively associated with levels of proximity, and specifically gestures associated with affiliation (greeting, gesture to mutually groom) were related to proximity. Synchronized low-intensity pant-hoots were also positively related to proximity in pairs of chimpanzees. Further, there were differences in the size of individual chimpanzees' proximity networks-the number of social relationships they maintained with others. Focal chimpanzees with larger proximity networks had a higher rate of both synchronized low- intensity pant-hoots and synchronized high-intensity pant-hoots. These results suggest that in addition to grooming, both gestures and synchronized vocalizations may play key roles in allowing chimpanzees to manage a large and differentiated set of social relationships. Gestures may be important in reducing the aggression arising from being in close proximity to others, allowing for proximity to be maintained for longer and facilitating grooming. Vocalizations may allow chimpanzees to communicate with a larger number of recipients than gestures and the synchronized nature of the pant-hoot calls may facilitate social bonding of more numerous social relationships. As group sizes increased through human evolution, both gestures and synchronized vocalizations may have played important roles in bonding social relationships in a more time-efficient manner than grooming.