Chimpanzee Communities Research Articles

Dialects in leaf-clipping and other leaf-modifying gestures between neighbouring communities of East African chimpanzees

Dialects are a cultural property of animal communication previously described in the signals of several animal species. While dialects have predominantly been described in vocal signals, chimpanzee leaf-clipping and other ‘leaf-modifying’ gestures, used across chimpanzee and bonobo communities, have been suggested as a candidate for cultural variation in gestural communication. Here we combine direct observation with archaeological techniques to compare the form and use of leaf-modifying gestures in two neighbouring communities of East African chimpanzees. We found that while both communities used multiple forms, primarily within sexual solicitation, they showed a strong preference for a single, different gesture form. The observed variation in form preference between these neighbouring communities within the same context suggests that these differences are, at least in part, socially derived. Our results highlight an unexplored source of variation and flexibility in gestural communication, opening the door for future research to explore socially derived dialects in non-vocal communication.

Association patterns in a high-elevation chimpanzee community in Rwanda

Investment in social interaction and affiliative behaviour is often related to variation in sex-specific dispersal patterns among species but can also vary within species in response to local environmental conditions and feeding competition. Chimpanzee (Pan troglodytes) association patterns have been studied primarily in low and mid-elevation tropical forests. This study adds to our understanding of association patterns in chimpanzees by presenting the first data on association patterns in a high-elevation population at Nyungwe in equatorial Rwanda. Using data on co-occurrence in social parties (party association) and calculating gregariousness indices, we quantified levels of associations among age-sex classes relative to food (fruit) distribution and the presence of oestrous females. Overall, adult females in Nyungwe are less gregarious than males. Female but not male gregariousness increased when feeding on clumped foods, indicating that resource characteristics impact female association tendencies more intensely than males. Despite high elevation forests representing a habitat with relatively fewer fruit resources for chimpanzees, at an edge of range location, we found that gregariousness patterns were driven by similar factors as those in mid and low elevation chimpanzee populations.

Wild chimpanzee behavior suggests that a savanna-mosaic habitat did not support the emergence of hominin terrestrial bipedalism

Bipedalism, a defining feature of the human lineage, is thought to have evolved as forests retreated in the late Miocene-Pliocene. Chimpanzees living in analogous habitats to early hominins offer a unique opportunity to investigate the ecological drivers of bipedalism that cannot be addressed via the fossil record alone. We investigated positional behavior and terrestriality in a savanna-mosaic community of chimpanzees (Pan troglodytes schweinfurthii) in the Issa Valley, Tanzania as the first test in a living ape of the hypothesis that wooded, savanna habitats were a catalyst for terrestrial bipedalism. Contrary to widely accepted hypotheses of increased terrestriality selecting for habitual bipedalism, results indicate that trees remained an essential component of the hominin adaptive niche, with bipedalism evolving in an arboreal context, likely driven by foraging strategy.

Repellent activity against Anopheles gambiae of the leaves of nesting trees in the Sebitoli chimpanzee community of Kibale National Park, Uganda.

BackgroundEvery evening, chimpanzees (Pan troglodytes) build a sleeping platform so called “nest” by intertwining branches of tree. Most of chimpanzees’ communities studied have a preference for tree species in which they nest. As female mosquitoes are feeding on the blood of their host at nighttime, chimpanzees may prevent being disturbed and bitten by mosquitoes by selecting tree species having properties to repel them.MethodsTo test the hypothesis that chimpanzees choose tree species for their aromatic properties, data related to 1,081 nesting trees built between 2017 and 2019 in the Sebitoli community of Kibale National Park (Uganda) were analysed. The 10 most used trees were compared to the 10 most common trees in the habitat that were not preferred for nesting. Leaves from the 20 trees species were collected and hydro-distillated to obtain essential oils and one of the by-products for behavioural bioassays against females of the African mosquito, Anopheles gambiae.ResultsSebitoli chimpanzees showed tree preferences: 10 species correspond to more than 80% of the nesting trees. Out of the essential oil obtained from the 10 nesting trees, 7 extracts for at least one concentration tested showed spatial repellency, 7 were irritant by contact and none were toxic. In the other hand, for the abundant trees in their habitat not used by chimpanzees, only 3 were repellent and 5 irritants.Discussion and conclusionThis study contributes to evidence that chimpanzees, to avoid annoying mosquitoes, may select their nesting trees according to their repellent properties (linked to chemical parameters), a potential inspiration for human health.Supplementary InformationThe online version contains supplementary material available at 10.1186/s12936-022-04291-7.

Leader election of dynamic wireless intelligent control machine in sensor network distributed processing

Strategies to streamline energy resources in wireless sensor networks (WSN) are still a hot topic. Many researchers have proposed various models of the WSN protocol so that the base station (BS)/sink can receive data collected from each sensor node. The energy in the WSN, which has a long life span, will convey a lot of data to the sink until the sensor nodes die. This paper has proposed modeling the WSN protocol by utilizing the classification and clustering of sensor nodes using the chimpanzee leader election optimization (CLEO) method. The CLEO algorithm is a meta-heuristic algorithm inspired by a social model that applies the pattern of leader selection to the chimpanzee community. The optimization method has the advantage of getting a fast fitness value which is one of the critical factors in utilizing meta-heuristic models in the WSN field. The closest sensor nodes at a certain distance to the sink are marked as classification, while other sensor nodes are marked as clusters. The clustering of sensor nodes utilizes a cluster function using the CLEO algorithm. This paper also carried out the selection of group leaders to obtain an energy efficiency model from WSN-CLEO. The results of the proposed method, WSN-CLEO, have been compared to other protocol models.

Flexibility in the social structure of male chimpanzees (Pan troglodytes schweinfurthii) in the Budongo Forest, Uganda.

Individuals of social species experience competitive costs and social benefits of group living. Substantial flexibility in humans' social structure and the combination of different types of social structure with fission–fusion dynamics allow us to live in extremely large groups—overcoming some of the costs of group living while capitalizing on the benefits. Non-human species also show a range of social strategies to deal with this trade-off. Chimpanzees are an archetypical fission–fusion species, using dynamic changes in day-to-day association to moderate the costs of within-group competition. Using 4 years of association data from two neighbouring communities of East African chimpanzees (Pan troglodytes schweinfurthii), we describe an unexplored level of flexibility in chimpanzee social structure. We show that males from the larger Waibira community (N = 24–31) exhibited additional structural levels of semi-stable core–periphery society, while males from the smaller Sonso community (N = 10–13) did not. This novel core–periphery pattern adds to previous results describing alternative modular social structure in other large communities of chimpanzees. Our data support the hypothesis that chimpanzees can incorporate a range of strategies in addition to fission–fusion to overcome costs of social living, and that their social structures may be closer to that of modern humans than previously described.

Adaptive Party Choice of Low-Ranking Males in Fission-Fusion Dynamics of Chimpanzees in Kalinzu Forest Reserve, Uganda.

Simple SummaryVarious mammalian species, including primates, exhibit fission–fusion dynamics (FFD), which alters the size of subgroups through splitting and merging. Among primates, chimpanzees (Pan troglodytes) have been particularly studied as a species that exhibits a high degree of FFD, wherein members of the same group form temporary subgroups (parties) that vary in both size and composition. Most studies on the FFD of chimpanzees have focused on its role in reducing food competition among individuals. In this study, we focused on the association patterns of each adult male chimpanzee and examined whether the social factors, including male dominance rank and aggression from other males, affect their party attendance behavior. Low-ranking males spent more time alone than other males when mating opportunities were absent. When females were available for mating, males of all ranks showed a similar party attendance behavior. The aggression increased with the number of males in the party, and low-ranking males received more aggression than higher-ranking males. These results suggest that low-ranking males frequently traveled alone to avoid aggression from other males unless they attended parties to seek mating opportunities. The FFD seemed to offer alternative tactics for low-ranking males to balance the costs and benefits of attending parties.Several studies have examined factors that regulate fission–fusion dynamics (FFD) in chimpanzee communities, such as receptive females, predation risks, and food availability. However, the effects of these factors vary between populations. In this study, we conducted focal animal observations of adult males in the M group in Kalinzu to examine the influence of male dominance rank, aggression from other males, the presence of females exhibiting maximum sexual swelling (MS), and fruit abundance on male tendencies of party attendance. We found that low-ranking males spent more time alone than other males when females with MS were absent. In contrast, when females with MS were present, males of all ranks showed a similar tendency of party attendance. We also found that the aggressive interactions increased with the number of males irrespective of the presence or absence of females with MS, and low-ranking males attracted aggression more frequently than higher-ranking males. These results suggest that low-ranking males frequently ranged alone to avoid aggression from other males unless they attended parties to seek mating opportunities. We conclude that low-ranking males have alternative tactics to balance the costs and benefits incurred or gained when attending parties.

Population-specific call order in chimpanzee greeting vocal sequences

SummaryPrimates rarely learn new vocalizations, but they can learn to use their vocalizations in different contexts. Such “vocal usage learning,” particularly in vocal sequences, is a hallmark of human language, but remains understudied in non-human primates. We assess usage learning in four wild chimpanzee communities of Taï and Budongo Forests by investigating population differences in call ordering of a greeting vocal sequence. Whilst in all groups, these sequences consisted of pant-hoots (long-distance contact call) and pant-grunts (short-distance submissive call), the order of the two calls differed across populations. Taï chimpanzees consistently commenced greetings with pant-hoots, whereas Budongo chimpanzees started with pant-grunts. We discuss different hypotheses to explain this pattern and conclude that higher intra-group aggression in Budongo may have led to a local pattern of individuals signaling submission first. This highlights how within-species variation in social dynamics may lead to flexibility in call order production, possibly acquired via usage learning.

Larger territories reduce mortality risk for chimpanzees, wolves, and agents: Multiple lines of evidence in a model validation framework

Validation of agent-based models is underappreciated in scientific studies even though this process is an important part of ensuring that a model is a reliable research tool. Here we propose modifications to a model validation framework and illustrate this framework with a case study of territorial behavior. In species that defend territories, larger territories provide obvious benefits, such as increased access to food, shelter, and mates. An additional potential benefit is that larger territories could provide protection from intergroup conflict. Considerations from geometry indicate that per capita risk of death from intergroup violence should decrease with increasing territory size, insofar as conflict occurs mainly at the periphery. We tested this inference using computer simulations and data from chimpanzees (Pan troglodytes) and gray wolves (Canis lupus). We designed an agent-based model that allows territories to vary freely in size and shape. We present a framework for model validation and apply it to our agent-based model to substantiate its accuracy. Simulations from the validated model confirmed the predictions from geometry. Under a broad range of parameter values, per capita mortality rate decreased in larger territories. Similarly, using published data on rates of death from intercommunity aggression in 16 chimpanzee communities, as well as new data from 38 wolf packs, we found that per capita mortality rate correlated negatively with a measure of territory size. These findings indicate that in species with lethal intergroup aggression, one simple aspect of territory geometry – size – has strong effects on mortality. Several lines of evidence indicate that the per capita rates of mortality from warfare have decreased in many human societies over time. One factor contributing to this decrease may be the increasing geographic extent of political entities.

Characterization of Pan social systems reveals in-group/out-group distinction and out-group tolerance in bonobos

Human between-group interactions are highly variable, ranging from violent to tolerant and affiliative. Tolerance between groups is linked to our unique capacity for large-scale cooperation and cumulative culture, but its evolutionary origins are understudied. In chimpanzees, one of our closest living relatives, predominantly hostile between-group interactions impede cooperation and information flow across groups. In contrast, in our other closest living relative, the bonobo, tolerant between-group associations are observed. However, as these associations can be frequent and prolonged and involve social interactions that mirror those within groups, it is unclear whether these bonobos really do belong to separate groups. Alternatively, the bonobo grouping patterns may be homologous to observations from the large Ngogo chimpanzee community, where individuals form within-group neighborhoods despite sharing the same membership in the larger group. To characterize bonobo grouping patterns, we compare the social structure of the Kokolopori bonobos with the chimpanzee group of Ngogo. Using cluster analysis, we find temporally stable clusters only in bonobos. Despite the large spatial overlap and frequent interactions between the bonobo clusters, we identified significant association preference within but not between clusters and a unique space use of each cluster. Although bonobo associations are flexible (i.e., fission-fusion dynamics), cluster membership predicted the bonobo fission compositions and the spatial cohesion of individuals during encounters. These findings suggest the presence of a social system that combines clear in-group/out-group distinction and out-group tolerance in bonobos, offering a unique referential model for the evolution of tolerant between-group interactions in humans.

Well-digging in a community of forest-living wild East African chimpanzees (Pan troglodytes schweinfurthii)

Access to resources shapes species’ physiology and behaviour. Water is not typically considered a limiting resource for rainforest-living chimpanzees; however, several savannah and savannah-woodland communities show behavioural adaptations to limited water. Here, we provide a first report of habitual well-digging in a rainforest-living group of East African chimpanzees (Pan troglodytes schweinfurthii) and suggest that it may have been imported into the community’s behavioural repertoire by an immigrant female. We describe the presence and frequency of well-digging and related behaviour, and suggest that its subsequent spread in the group may have involved some degree of social learning. We highlight that subsurface water is a concealed resource, and that the limited spread of well-digging in the group may highlight the cognitive, rather than physical, challenges it presents in a rainforest environment.

Chimpanzee leader election optimization

Solving the convergence problem in the world of computing is needed to find an optimization solution. Many researchers propose a meta-heuristic approach in solving problems quickly using the results of the approach. This method is used because of its flexibility and fast search rate. Each meta-heuristic method has its own characteristics and is not necessarily able to solve the problem for all cases. This paper proposes a meta-heuristic method that is inspired by the way in which leaders are chosen in the chimpanzee community. The leader in the chimpanzee community is always the alpha male chimpanzee and has relationships with all chimps, both male and female. The principle of Chimpanzee Leader Election Optimization (CLEO) is the phase of improving performance and social relations with community members. The chimp performance improvement phase is based on the age aspect, which affects the memorize’s ability of the chimps, which are inverted U-shaped. Meanwhile, the social relations/mating phase has been analogized in this CLEO algorithm. This paper has also evaluated the performance of the proposed algorithm, both to find the optimal value in the minimum and maximum search. This algorithm has also been compared with various other meta-heuristic algorithms. The results of the CLEO algorithm are quite competitive with other meta-heuristic algorithms in multi-high dimensional.

Referential gestures are not ubiquitous in wild chimpanzees: alternative functions for exaggerated loud scratch gestures

A fundamental aspect of human communication is our ability to refer to external objects and events through both words and gestures (such as pointing), yet the evolutionary origins of such signals remain obscure. Apes, living in their natural environments, rarely or never point, but it has been claimed that male chimpanzees, Pan troglodytes schweinfurthii, from the Ngogo community, Uganda, habitually use exaggerated loud scratches (ELSs) to refer to specific body locations where they wish to be groomed (Pika & Mitani, 2006, Current Biology, 16(6), 191–192). This study suggested continuity between referential abilities in humans and our closest living relatives, making it an important finding to replicate in other populations. Hence here, we compared whether ELSs are used in a referential manner across four wild communities of eastern chimpanzees (Ngogo, Kanyawara, Sonso and Waibira). Our data show that scratchers were significantly more likely to receive grooming in the scratched location at Ngogo compared to the other three sites. At the latter sites this response occurred at low rates and signallers did not seem to pursue this goal. This suggests that ELSs do not function referentially at these sites, and the published findings from Ngogo were not replicated. Further exploration into alternative functions of ELSs in the Kanyawara community revealed that, in this community, this signal functions to initiate grooming bouts and to reengage partners during grooming pauses. Individuals who produced the signal to initiate grooming were likely to offer grooming. In contrast, during grooming bouts, groomers produced ELSs to request reciprocation of grooming from their partner. Our study demonstrates that chimpanzees do not ubiquitously use the ELS in a referential manner, but that they can use this gesture in a highly flexible fashion, with signal function depending on the intricate details of the social contexts in which they are produced.

Investigating foundations for hominin fire exploitation: Savanna-dwelling chimpanzees (Pan troglodytes verus) in fire-altered landscapes

Humans' extensive use of fire is one behavior that sets us apart from all other animals. However, our ancestors' reliance on controlled forms of fire—i.e., for cooking—was likely preceded by a long familiarity with fire beginning with passive exploitation of naturally burned landscapes and followed by intermediate steps including active ecological modification via intentional burning. Here we explore our pyrophilic beginnings using observational data from savanna-dwelling chimpanzees. These data highlight the extent to which anthropogenic burning impacts the behavior and ecology of sympatric primates and provides an opportunity to study the ways in which apes living in a fire-altered world exploit opportunities presented by burning. Using monthly burn scar data and daily range use data we quantify the impact of burning episodes on chimpanzee habitat. Over the course of one dry season, approximately 74% of the total estimated range of the Fongoli community of savanna-dwelling chimpanzees (Pan troglodytes verus) was impacted by fire. We combine fire occurrences with behavioral data to test for relationships between burning and rate of encounter with food items and duration of subsequent patch residence time. Results show more frequent encounters and shorter patch residence times in burned areas. These data can be leveraged as a frame of reference for conceptualizing our extinct relatives’ behavior around fire.

All-You-Can-Eat: Influence of Proximity to Maize Gardens on the Wild Diet and the Forest Activities of the Sebitoli Chimpanzee Community in Kibale National Park.

Simple SummaryUnderstanding the resilience of primate populations to the threat of agricultural expansion is critical for effective conservation. Based on individual monitoring from morning to evening of wild chimpanzees in and around a protected area, we showed that the availability of maize at the forest edge had little effect on their activity budget by less resting and no impact on their wild diet and energy expenditure. In this area, large, caloric wild fruits are available year-round, and we observed no behavioral or dietary changes regarding wild resource availability either. Thus, the chimpanzees consume maize opportunistically as a bonus treat in their diet, and the presence of this nutritious resource does not seem to affect their role in seed dispersal and forest regeneration.Frugivorous primates have developed several strategies to deal with wild fruit scarcity, such as modifying their activity budget or enlarging their diet. Agricultural expansion threatens primate habitats and populations (e.g., disease transmission, agrochemical exposure), but it also increases crop feeding opportunities. We aimed at understanding whether maize presence close to the natural habitat of chimpanzees, a threatened species, would lead to significant behavioral modifications. We monitored 20 chimpanzees over 37 months in Kibale National Park, Uganda, with maize gardens at the forest edge. Based on focal nest-to-nest data, we analyzed their diet, activity budget, and energy balance depending on wild fruit and maize availability. We found that the Sebitoli area is a highly nutritive habitat for chimpanzees, with large and caloric wild fruits available all year long. The chimpanzees opportunistically consume maize and exploit it by resting less during maize season. However, no significant variation was found in daily paths and energy expenditures according to maize availability. No behavioral or energy modification was observed regarding wild resources either. Despite the availability of nutritious domestic resources, chimpanzees still exploit wild fruits and do not limit their movements. Thus, their contribution to seed dispersal and forest regeneration in this area is not affected.

<Note>Indirect evidence of tool-assisted hunting in the Bantan chimpanzee (Pan troglodytes verus) community in southeastern Senegal savanna-woodlands

Chimpanzees use tools extensively, but tool-assisted hunting has been reported at only two sites thus far. Systematic tool-assisted hunting has only been recorded at the Fongoli site in Senegal, where hundreds of cases have been recorded of chimpanzees using tools to hunt bushbaby (Galago senegalensis) prey. Here, we report a putative case of tool-assisted hunting at the Bantan study site in southeastern Senegal, based on indirect evidence of tools and tool-making traces in a context similar to that observed at neighboring Fongoli chimpanzee community. If our interpretations are correct, the simplest explanation is that Fongoli females have dispersed to this neighboring community, bringing with them the behavior of tool-assisted hunting. However, the persistence and frequency of tool-assisted hunting at other sites in Senegal is still unknown. Tool-assisted hunting in the Bantan chimpanzee community would present only the third such known site for such behavior among wild chimpanzees thus far.

Inter-community behavioural variation confirmed through indirect methods in four neighbouring chimpanzee communities in Cantanhez NP, Guinea-Bissau.

Culture, while long viewed as exclusively human, has now been demonstrated across diverse taxa and contexts. However, most animal culture data are constrained to well-studied, habituated groups. This is the case for chimpanzees, arguably the most ‘cultural’ non-human species. While much progress has been made charting wild chimpanzees' cultural repertoire, large gaps remain in our knowledge of the majority of the continent's chimpanzees. Furthermore, few studies have compared neighbouring communities, despite such comparisons providing the strongest evidence for culture, and few have studied communities living in anthropogenic habitats although their culture is in imminent danger of disappearing. Here we combine direct, indirect and remote methods, including camera traps, to study, over 2 years, four unhabituated neighbouring chimpanzee communities inhabiting human-impacted habitats in Cantanhez NP, Guinea-Bissau. From traces collected during 1089 km of reconnaissance walks and 4197 videos from 56 camera trap locations, we identified 18 putative cultural traits. These included some noteworthy novel behaviours for these communities, and behaviours possibly new to the species. We created preliminary behavioural profiles for each community, and found inter-community differences spanning tool use, communication, and social behaviour, demonstrating the importance of comparing neighbouring communities and of studying previously neglected communities including those inhabiting anthropogenic landscapes.

Collective knowledge and the dynamics of culture in chimpanzees.

Social learning in non-human primates has been studied experimentally for over 120 years, yet until the present century this was limited to what one individual learns from a single other. Evidence of group-wide traditions in the wild then highlighted the collective context for social learning, and broader ‘diffusion experiments’ have since demonstrated transmission at the community level. In the present article, we describe and set in comparative perspective three strands of our recent research that further explore the collective dimensions of culture and cumulative culture in chimpanzees. First, exposing small communities of chimpanzees to contexts incorporating increasingly challenging, but more rewarding tool use opportunities revealed solutions arising through the combination of different individuals' discoveries, spreading to become shared innovations. The second series of experiments yielded evidence of conformist changes from habitual techniques to alternatives displayed by a unanimous majority of others but implicating a form of quorum decision-making. Third, we found that between-group differences in social tolerance were associated with differential success in developing more complex tool use to exploit an increasingly inaccessible resource. We discuss the implications of this array of findings in the wider context of related studies of humans, other primates and non-primate species.This article is part of a discussion meeting issue ‘The emergence of collective knowledge and cumulative culture in animals, humans and machines’.

Overlooked evidence for semantic compositionality and signal reduction in wild chimpanzees (Pan troglodytes)

Recent discoveries of semantic compositionality in Japanese tits have enlivened the discussions on the presence of this phenomenon in wild animal communication. Data on semantic compositionality in wild apes are lacking, even though language experiments with captive apes have demonstrated they are capable of semantic compositionality. In this paper, I revisit the study by Boesch (Hum. Evol. 6:81–89, 1991) who investigated drumming sequences by an alpha male in a chimpanzee (Pan troglodytes) community in the Taï National Park, Côte d’Ivoire. A reanalysis of the data reveals that the alpha male produced semantically compositional combined messages of travel direction change and resting period initiation. Unlike the Japanese tits, the elements of the compositional expression were not simply juxtaposed but displayed structural reduction, while one of the two elements in the expression coded the meanings of both elements. These processes show relative resemblance to blending and fusion in human languages. Also unlike the tits, the elements of the compositional expression did not have a fixed order, although there was a fixed distribution of drumming events across the trees used for drumming. Because the elements of the expression appear to carry verb-like meanings, the compositional expression also resembles simple verb-verb constructions and short paratactic combinations of two clauses found across languages. In conclusion, the reanalysis suggests that semantic compositionality and phenomena resembling paratactic combinations of two clauses might have been present in the communication of the last common ancestor of chimpanzees and humans, not necessarily in the vocal modality.

Exploring greetings and leave-takings: communication during arrivals and departures by chimpanzees of the Bossou community, Guinea.

In human fission-fusion societies, ritualized non-linguistic signal exchanges that include gestures, vocalizations, and facial expressions are regularly observed at both arrivals (greetings) and departures (leave-takings). These communicative events play an important role in the formation and maintenance of social relationships. Wild chimpanzees also form large communities that split into smaller fluid parties during daily activities, with individuals moving freely between them. However, in chimpanzees only greetings have been reported. This study explores signal exchanges in the Bossou chimpanzee community during fissions (departures) and fusions (arrivals) given an individual's social rank, kinship, position as traveller or party-member, the level of potential threat, and the party size and presence of mature males. We analysed three time periods (1993-1994; 2003-2004; 2013-2014) during which the composition and social hierarchy of the community varied. We show that the occurrence and form of communication during fission and fusion events are mediated by social factors, including rank, kinship, and party size and composition. Individuals were more likely to communicate during fusions than during fissions, communication was more likely to be produced towards a higher-ranking individual and to non-kin individuals, but the tendency to communicate in general increased with an increase in social rank. The presence of more individuals, and in particular mature males, decreased the likelihood of communication. Communication during fusions supported patterns reported in previous studies on greetings, and our results support the argument that, if present, leave-takings are not a common feature of chimpanzee social interactions. Current methodological difficulties regarding the function of declarative signals hinder our ability to discriminate potential parting rituals within communication before departures. Given similar methodological difficulties, we also provide a note of caution in the interpretation of all signals produced during fusions as 'greetings'.