Chimpanzees communicate to coordinate a cultural practice.

Human culture thrives by virtue of communication, yet whether communication plays an influential role in the cultural lives of other animals remains understudied. Here, we investigated whether chimpanzees use communication to engage in a cultural practice by analysing grooming handclasp (GHC) interactions-a socio-cultural behaviour requiring interindividual coordination for successful execution. Previous accounts attributed GHC initiations to behavioural shaping, whereby the initiator physically moulds the partner's arm into the desired GHC posture. Using frame-by-frame analysis and matched-control methodology, we find that chimpanzees do not only shape their partner's posture (22%), but also use gestural communication to initiate GHC (44%), which requires an active and synchronized response from the partner. Moreover, in a third (34%) of the GHC initiations, the requisite coordination was achieved by seemingly effortless synchrony. Lastly, using a longitudinal approach, we find that for GHC initiations, communication occurs more frequently than shaping in experienced dyads and less in mother-offspring dyads. These findings are consistent with ontogenetic ritualization, thereby reflecting first documentation of chimpanzees communicating to coordinate a cultural practice. We conclude that chimpanzees show interactional flexibility in the socio-cultural domain, opening the possibility that the interplay between communication and culture is rooted in our deep evolutionary history.

Intentional gestural communication amongst red-capped mangabeys (Cercocebus torquatus).

Apes, human's closest living relatives, are renowned for their intentional and highly flexible use of gestural communication. In stark contrast, evidence for flexible and intentional gestural communication in monkeys is scarce. Here, we investigated the intentionality and flexibility of spontaneous gesture use in red-capped mangabeys (Cercocebus torquatus). We applied established methods used in ape gesture research to analyse whether the body acts produced by a total of 17 individuals living in three different groups in captivity qualified as intentionally produced gesture instances. Results showed that signallers showed all hallmarks of intentionality during the production of 20 out of a total of 21 different types of body acts. These were only produced in the presence of other individuals, and the monkeys showed audience checking, sensitivity to the attentional states of recipients, adjustment of signal modality, and response waiting relative to their production. Moreover, in case of communication failure, the monkeys showed goal persistence, and regarding the production contexts they showed some signs of means-ends dissociation. Therefore, these monkeys are capable of flexible and intentional gestural communication and use this to communicate with conspecifics. Our results corroborate recent findings showing that intentional gestural communication was already present in the monkey lineage of catarrhine primates. We discuss our results in light of the comparative approach towards human language evolution and highlight our finding that these monkeys also showed flexible and intentional use of four 'free' manual gesture types.

Mineral Acquisition from Clay by Budongo Forest Chimpanzees.

Chimpanzees of the Sonso community, Budongo Forest, Uganda were observed eating clay and drinking clay-water from waterholes. We show that clay, clay-rich water, and clay obtained with leaf sponges, provide a range of minerals in different concentrations. The presence of aluminium in the clay consumed indicates that it takes the form of kaolinite. We discuss the contribution of clay geophagy to the mineral intake of the Sonso chimpanzees and show that clay eaten using leaf sponges is particularly rich in minerals. We show that termite mound soil, also regularly consumed, is rich in minerals. We discuss the frequency of clay and termite soil geophagy in the context of the disappearance from Budongo Forest of a formerly rich source of minerals, the decaying pith of Raphia farinifera palms.

Social, contextual, and individual factors affecting the occurrence and acoustic structure of drumming bouts in wild chimpanzees (Pan troglodytes).

The production of structured and repetitive sounds by striking objects is a behavior found not only in humans, but also in a variety of animal species, including chimpanzees (Pan troglodytes). In this study we examined individual and social factors that may influence the frequency with which individuals engage in drumming behavior when producing long distance pant hoot vocalizations, and analyzed the temporal structure of those drumming bouts. Male chimpanzees from Budongo Forest, Uganda, drummed significantly more frequently during travel than feeding or resting and older individuals were significantly more likely to produce drumming bouts than younger ones. In contrast, we found no evidence that the presence of estrus females, high ranking males and preferred social partners in the caller's vicinty had an effect on the frequency with which an individual accompanied their pant hoot vocalization with drumming. Through acoustic analyses, we demonstrated that drumming sequences produced with pant hoots may have contained information on individual identity and that qualitatively, there was individual variation in the complexity of the temporal patterns produced. We conclude that drumming patterns may act as individually distinctive long-distance signals that, together with pant hoot vocalizations, function to coordinate the movement and spacing of dispersed individuals within a community, rather than as signals to group members in the immediate audience.

'Adoption' by maternal siblings in wild chimpanzees.

The adoption of unrelated orphaned infants is something chimpanzees and humans have in common. Providing parental care has fitness implications for both the adopter and orphan, and cases of adoption have thus been cited as evidence for a shared origin of an altruistic behaviour. We provide new data on adoptions in the free-living Sonso chimpanzee community in Uganda, together with an analysis of published data from other long-term field sites. As a default pattern, we find that orphan chimpanzees do not become adopted by adult group members but wherever possible associate with each other, usually as maternal sibling pairs. This occurs even if both partners are still immature, with older individuals effectively becoming 'child household heads'. Adoption of orphans by unrelated individuals does occur but usually only if no maternal siblings or other relatives are present and only after significant delays. In conclusion, following the loss of their mother, orphaned chimpanzees preferentially associate along pre-existing social bonds, which are typically strongest amongst maternal siblings.

Chimpanzee alarm call production meets key criteria for intentionality.

Determining the intentionality of primate communication is critical to understanding the evolution of human language. Although intentional signalling has been claimed for some great ape gestural signals, comparable evidence is currently lacking for their vocal signals. We presented wild chimpanzees with a python model and found that two of three alarm call types exhibited characteristics previously used to argue for intentionality in gestural communication. These alarm calls were: (i) socially directed and given to the arrival of friends, (ii) associated with visual monitoring of the audience and gaze alternations, and (iii) goal directed, as calling only stopped when recipients were safe from the predator. Our results demonstrate that certain vocalisations of our closest living relatives qualify as intentional signals, in a directly comparable way to many great ape gestures. We conclude that our results undermine a central argument of gestural theories of language evolution and instead support a multimodal origin of human language.

Network analysis of social changes in a captive chimpanzee community following the successful integration of two adult groups.

Chimpanzees are highly territorial and have the potential to be extremely aggressive toward unfamiliar individuals. In the wild, transfer between groups is almost exclusively completed by nulliparous females, yet in captivity there is often a need to introduce and integrate a range of individuals, including adult males. We describe the process of successfully integrating two groups of chimpanzees, each containing 11 individuals, in the Budongo Trail facility at the Royal Zoological Society of Scotland's Edinburgh Zoo. We use social network analysis to document changes in group dynamics within this population over the 16 months following integration. Aggression rates were low overall and members of the two original groups engaged in significantly fewer aggressive interactions over time. Association and grooming data indicate that relationships between members of the original groups became stronger and more affiliative with time. Despite these positive indicators the association data revealed the continued existence of two distinct subgroups, a year after integration. Our data show that when given complex space and freedom to exhibit natural fission-fusion groupings, in which the chimpanzees choose whom they wish to associate and interact with, the building of strong affiliative relationships with unfamiliar individuals is a very gradual process.

Predator and non-predator long-distance calls in Guereza colobus monkeys.

East African Guereza colobus monkey males are known for their conspicuous roaring behaviour; a spectacle that can dominate the predawn hours of African forests. Recent research has shown that these monkeys also produce roars during daytime hours in response to predators. While roars to leopards and eagles differ in how roaring phrases are assembled into sequences, there are no obvious structural differences between predawn roars and roars to eagles. Although recipients could use daytime information to disambiguate between the two contexts, this may be a risky strategy because eagles can be active before dawn. We carried out acoustic analyses, which showed that the duration of the first roaring phrase was significantly longer in predawn roars compared to eagle roars. Furthermore, the initial call repetition rate was faster in response to eagle roars compared to predawn roars. Apart from these two differences, all other acoustic characteristics were identical between the two contexts. Although these monkeys exhibit some of the most basic vocal behaviour found in non-human primates, callers are able to provide reliable contextual information by varying the duration and assemblage of individual vocal units. Playback experiments are needed to confirm whether recipients relate these acoustic differences to different contexts.

The alarm call system of two species of black-and-white colobus monkeys (Colobus polykomos and Colobus guereza).

Vervet monkey alarm calling has long been the paradigmatic example of how primates use vocalizations in response to predators. In vervets, there is a close and direct relationship between the production of distinct alarm vocalizations and the presence of distinct predator types. Recent fieldwork has however revealed the use of several additional alarm calling systems in primates. Here, the authors describe playback studies on the alarm call system of two colobine species, the King colobus (Colobus polykomos) of Taï Forest, Ivory Coast, and the Guereza colobus (C. guereza) of Budongo Forest, Uganda. Both species produce two basic alarm call types, snorts and acoustically variable roaring phrases, when confronted with leopards or crowned eagles. Neither call type is given exclusively to one predator, but the authors found strong regularities in call sequencing. Leopards typically elicited sequences consisting of a snort followed by few phrases, while eagles typically elicited sequences with no snorts and many phrases. The authors discuss how these call sequences have the potential to encode information at different levels, such as predator type, response-urgency, or the caller's imminent behavior.

Male blue monkeys alarm call in response to danger experienced by others.

Male blue monkeys (Cercopithecus mitis stuhlmanni) of Budongo Forest, Uganda, produce two acoustically distinct alarm calls: hacks to crowned eagles (Stephanoaetus coronatus) and pyows to leopards (Panthera pardus) and a range of other disturbances. In playback experiments, males responded to leopard growls exclusively with a series of pyows and to eagle shrieks predominantly with hacks. Responses to playbacks of these alarm call series matched the responses to the corresponding predators, suggesting that the calls conveyed something about the nature of the threat. When responding to a series of hacks, indicating an eagle, males responded predominately with hacks, but produced significantly more calls if their group members were close to the playback stimulus than far away, regardless of their own position. When responding to a series of pyows, indicating a range of disturbances, males responded with pyows, but call rates were independent of the distance of other group members. The results suggest that males took into account the degree of danger experienced by other group members.

Geographic variation in Thomas langur (Presbytis thomasi) loud calls.

Geographic variation in primate vocalizations has been described at two levels. First, at the level of acoustic variation within the same call type between populations and, second, at the level of presence or absence of certain call types in different populations. Acoustic variation is of interest because there are several factors that can explain this variation, such as gene flow, ecological factors and population density. Here we focus on the first level in a Southeast Asian primate, the Thomas langur. We recorded male loud calls in four populations that differed in their geographic distances from each other and had varying geographic barriers in between them, such as rivers and mountain ranges. The presence of these barriers leads to expectations of loud call variation under the gene flow model, which are examined here. We conducted a principal components analysis to condense the number of acoustic variables. With a subsequent discriminant function analysis on the six principal component scores, we found that the percentage of loud calls that were correctly assigned to a population was relatively high (50.0-76.2%) when three randomly selected loud calls from each male were used. Using the discriminant functions from this analysis to predict population membership of the remainder of the loud calls yielded lower, but still relatively high correct assignment percentages (26.2-66.7%). Analyses to examine the influence of barriers on similarities between populations confirm our expectations. We discuss that differences in loud calls are probably most parsimoniously explained by gene flow (or the lack thereof) between the populations and that future studies of genetic differences are crucial to test this hypothesis.