Studies of nesting behaviours of free-ranging apes typically focus on ecological variables such as preferred tree species and areas within the home range, heights of nests, and nest group sizes. However, nesting in captive apes is rarely studied, despite the ubiquity of this sleep-related behaviour. The paucity of field data is often attributed to the inherent difficulty in observing what is essentially a nighttime behaviour. Captive settings can provide researchers with an ideal opportunity to record nesting and sleep-related behaviours, yet such research on captive apes is also scant. Topics addressed include current practices in zoos regarding conditions for sleep in great apes, the potential effects of social and environmental factors on sleep site selection, the motor patterns involved in nest construction, preferred nesting structures and substrates, and nocturnal behaviours. This thesis documented and empirically tested hypotheses concerning nest-related activities in captive chimpanzees, with an aim to generate practical recommendations for enclosure design, sleeping areas, sleeping structures, and nesting substrates that have implications for the welfare of captive apes. As with the few reports that already exist, most chimpanzees in this research frequently constructed night nests. When building a nest, some techniques appeared to be universal across individuals and groups, where others were group-specific or occasionally characteristic of only certain individuals. An experiment showed that specific materials are preferred over others for nest building. Many chimpanzees appeared to express persistent preferences for particular sleeping sites, and for some this was to maintain proximity to kin or other closely bonded individuals. In one group, individual sleeping site preferences changed across seasons, although again this was subject to individual differences. Video analyses of nighttime behaviours demonstrated that, although nests/sleep sites are primarily used for rest subsequent to retirement, a number of social and non-social activities were performed throughout the night. In conjunction with analysis of postural and orientation shifts, these data are unique in describing the nocturnal behaviours of chimpanzees out with a laboratory setting. Several aspects of nest-related behaviours showed a high degree of inter-and intra-group variation. Although this cautions against generalising findings across captive populations, research of this type has applied implications for the management of captive ape species, and can add to our as-yet meagre understanding of their nest and sleep-related behaviours.

No description available.

The concept of culture has recently been used to explain behavioural variation and trans-generational continuity of behaviour in non-human animals and in chimpanzees in particular. However, few studies in the wild have systematically investigated how the environment and behavioural adaptation might influence behavioural diversity. In this context, one habituated community of wild chimpanzees (Pan troglodytes verus) at Bossou, Guinea, and two neighbouring non-habituated communities in the Nimba Mountains region of Guinea and Cote d'Ivoire were the subject of a detailed study of behavioural variation at the intra- and inter-community level. An ecologically-based approach was adopted to investigate variation in nest building, in the use of the oil-palm tree (Elaeis guineensis), in ant-dipping and in tool-choice and -manufacturing. A significant influence of environmental variables on nesting parameters emerged explaining much of the variation observed between the three sites. However, some differences that arose are more likely to reflect differences in social structure and organisation. The comparative study of the utilisation of the oil-palm tree failed to reveal proximate environmental parameters that might explain significant observed variations in use. These findings raise interesting and important questions pertaining to diffusion of behaviour between neighbouring chimpanzee communities. Dipping for driver ants, Dorylus spp., is often cited as one of the best examples of culture in chimpanzees. A detailed analysis of this behaviour at Bossou suggests that risk exposure affects frequency of performance in the developing chimpanzee and reveals a strong influence of prey characteristics, including aggressiveness and/or gregariousness , on tool length and technique employed. Variations in tool-choice and tool-manufacturing within and between three tool-use behaviours at Bossoui nvolving the use of a stick or a stalk were found to be significantly associated with the nature of the task and its predictability, emphasising the importance of environmental affordance and constraints on these processes. In addition, efficiency in behaviour across another set of three tool-use behaviours was explored focusing chiefly on age-class differences. An analysis of individual and community-level patterns of laterality in hand-use between these three tool-use behaviours is also provided. The data supply some evidence to support the selective advantages of lateralization in hand-use with respect to behavioural efficiency. The findings also suggest that haptic tasks have played an important evolutionary role in driving population-level handedness, and reveal that although complex tool-uses exhibited high levels of lateralization, these failed to show task specialisation across individuals. Finally, this thesis presents a comprehensive review analysis of individual and community-wide variation across a range of behaviours observed in chimpanzees and identifies paths and hypotheses that warrant further exploration and testing with the aim to gain further insight into cultural processes in nonhuman animals.

Great ape gesture is an elaborate, flexible system of intentional communication. It has been suggested that human language originated in gesture, thus, the gestural communication of great apes is of great interest for questions on the origin of language. To date, systematic studies of great ape gesture have been limited to restricted captive settings, supplemented by the study of a few specific gestures in wild populations. To address questions about gestural communication from an evolutionary perspective it is necessary to extend the systematic study of gesture into a wild ape population. I therefore undertook a 22-month study of gesture in the wild Sonso chimpanzee community in Budongo, Uganda. Sonso chimpanzees employ a large repertoire of species-typical gestures in intentional communication; a proportion of this repertoire appears to be ape-typical, as would be expected with a biologically given trait. Chimpanzees can acquire new behavioural patterns through imitation; however, this apparently does not represent a significant means of acquiring gestures. Gesturing was employed regularly in an intentional manner from the end of the first year, and was used by chimpanzees of all ages to communicate across a range of contexts, including the evolutionarily urgent context of consortship. Immature chimpanzees used a wide range of gestures, which they combined into rapid sequences. With maturity, use of the repertoire was ‘tuned’ to focus on the most effective gestures, which were then used individually. Despite the evidence for referential pointing in captive chimpanzees, there was little evidence for the regular use of it in wild chimpanzees. Gestures were used to communicate a range of imperative requests that regulated social behaviour. Chimpanzee gestures vary from the ambiguous to the highly specific in meaning; and, while gestures were used flexibly, they tended to be associated with a single dominant meaning.

Evidence for conceptual semantics is well established in monkeys, however this basis of human language is less evident in the great apes. In order to study semantic communications in chimpanzees, I analyzed alarm calls produced towards a blimp as it was flying overhead. I then replayed a set of these alarm calls to the chimps on a different day. The chimps appeared to act in a manner consistent with the presence of the blimp. The calls they produced in response to the playback stimuli were nearly identical to the calls that were produced during the actual flyover. Though the data collected were not sufficient to support a definitive claim, it does appear that the chimpanzees of the study have a meaning-laden vocalization for the aerial stimuli. Whether this call is specific to the blimp or generalizable to other aerial threats is yet to be determined. / by Alyssa M. Raymond. / Thesis (M.A.)--Florida Atlantic University, 2012. / Includes bibliography. / Electronic reproduction. Boca Raton, Fla., 2012. Mode of access: World Wide Web.

In this thesis, I explore two subjects of importance to the study of cultural evolution and cumulative culture; behavioural flexibility in chimpanzees, and social transmission in human children. In Chapter 1, I give an overview of current literature on the cognitive requirements of cumulative culture, with a focus on behavioural flexibility as a capacity which facilitates cumulative culture. I also explore a current discussion in the field of cultural evolution; namely the debate between "standard" and cultural attraction-based approaches to the study of cultural evolution. Chapter 2 is an experimental investigation of the capacity of chimpanzees to respond flexibly to a changing foraging task. This study found that chimpanzees did alter their behaviour, but to a limited degree. In Chapter 3 I provide the same artificial foraging task to two further groups of chimpanzees, at a sanctuary in Zambia. This study again found that chimpanzees altered their behaviour in response to task constraints, but also found a significant difference in performance between the two groups tested. Chapter 4 explores one potential factor which may contribute to these group differences; social tolerance. Data on social tolerance from all three groups of chimpanzees is presented. In Chapter 5, I turn to another key factor in the study of culture and also address the cultural attraction approach, by conducting a transmission chain study of four- to eight-year-old human children, comparing the transmission of a symbolic and non-symbolic image. I found that neither image was reliably transmitted along transmission chains. Finally, in Chapter 6, I discuss the findings of the thesis, and suggest that future work considers multiple demographic groups, whether this means the inclusion of multiple groups of apes in studies of non-human primate cognition, or the consideration of how cultural behaviours might be transformed when transmitted by human children rather than adults.

An important element in understanding the evolutionary origin of human language is to explore homologous traits in cognition and communication between primates and humans (Burling, 1993, Hewes, 1973). One proposed modality of language evolution is that of gestural communication, defined as communicative movements of hands without using or touching objects (de Waal, 2003). While homologies between primate calls and language have been relatively well explored, we still have a limited understanding of how cognitive abilities may have shaped the characteristics of primate gestures (Corballis, 2003). Chimpanzees (Pan troglodytes) are our closest living relatives and display some complex cognitive skills in various aspects of their gestural behaviour in captivity (de Waal, 2003, Pollick and de Waal, 2007). However, it is not yet currently clear to what extent these abilities seen in captive apes are typical of chimpanzees in general and to what extent cognitive capacities observed in captive chimpanzees have been enhanced by the socio-cultural environment of captivity such as language training. In this Ph.D. research, I investigated the cognitive skills underlying gestural communication in both wild and language trained chimpanzees, with a special focus on the repertoire and the intentionality of production and comprehension. The study of cognitive skills underlying the production of the repertoire and the role of intentionality is important because these skills are cognitively demanding and are a prerequisite in human infants for their ability to acquire language (Baldwin, 1995, Olson, 1993). My research suggests that chimpanzee gestural communication is cognitively complex and may be homologous with the cognitive skills evident in pre-verbal infants on the cusp of language acquisition. Chimpanzees display a multifaceted and complex signal repertoire of manual gestures. These gestures are the prototypes, within which there is variation, and between which the boundaries are not clear-cut, but there is gradation apparent along several morphological components. Both wild and language trained chimpanzees communicate intentionally about their perceived desires and the actions that they want the recipients to undertake. They do not just express their emotions, but they communicate flexibly by adjusting their communicative tactics in response to the comprehension states of the recipient. Whilst chimpanzees communicate their intentions flexibly, the messages conveyed are specific. However, recipients comprehend gestures flexibly in light of the signaller’s overall intentions. Whilst wild and language trained chimpanzee gestural communication revealed similar cognitive characteristics, language trained chimpanzees outperformed wild apes in that they had ability to use signals which made distinctions that human deictic words can make. Whilst these differences between wild and language trained chimpanzees may be due to the different methodological approaches used, it is conceivable that language training may have influenced captive ape cognitive skills in the representational domain. These results from wild and language trained chimpanzees indicate that chimpanzees possess some form of cognitive skills necessary for language development and that cognitive skills underlying repertoire and use in chimpanzees are a shared capacity between humans, other apes and a common ancestor. These findings render theories of the gestural origins of language more plausible. Related publications: 1. Roberts, A. I., Vick, S.-J., Roberts, S. G. B., Buchanan-Smith, H. M. & Zuberbühler, K. 2012. A structure-based repertoire of manual gestures in wild chimpanzees: Statistical analyses of a graded communication system. Evolution and Human Behavior, Published online: http://dx.doi.org/ 10.1016/j.evolhumbehav.2012.05.006 2. Roberts, A. I., Vick, S.-J. & Buchanan-Smith, H. 2012. Usage and comprehension of manual gestures in wild chimpanzees. Animal Behaviour, Published online: http://dx.doi.org/10.1016/j.anbehav.2012.05.022

The following research paper was concerned with five basic objectives:(1) outlining the major theoretical and methodological approaches used in the reconstruction of early hominid social behavior/social structure as a context in which to view Richard Wrangham's and Michael Ghiglieri's phylogenetic models of stem-hominid social structure.(2) examining Wrangham's and Ghiglieri's models of stem-hominid and chimpanzee social structure.(3) indicating how theoretical and methodological aspects of structure essentially represent an extension of the theoretical and methodological approaches the same researchers applied to their models of chimpanzee social structure.(4) addressing the theoretical and methodological deficiences of Wrangham's and Ghiglieri's models of chimpanzee social structure.(5) providing suggestions for improved phylogenetic models of early hominid social structure.The first objective was achieved by: (a) reviewing Tooby and Devore's (1986) and Wrangham's (1986) evaluations of the major theoretical approaches and methodologies used in the reconstruction of hominid social behavior/structure (b) defining, classifying and evaluating Wrangham's and Ghiglieri's phylogenetic approaches within this context.The second objective was accomplished by outlining, analyzing, and comparing/contrasting Wrangham's and Ghiglieri's phylogenetic models of stem-hominid social structure (i.e.Wrangham 1986; Ghiglieri 1987, 1989) and Wrangham's and Ghiglieri's models of chimpanzee social structure (i.e. Wrangham 1975, 1979; Ghiglieri 1984, 1985, 1987, 1989).The third objective was achieved by recognizing how Wrangham and Ghiglieri used/stressed principles and concepts derived from evolutionary biology and/or behavioral ecology to develop their models of stem-hominid and chimpanzee social structure. This analysis showed that Wrangham's models of social structure were more favorably inclined toward the method of behavioral ecology than Ghiglieri's models, which favored a sociobiological paradigm. Furthermore, although neither researcher relied exclusively on the above theoretical approaches, the main thrust of their argument often centered around it. For instance, Wrangham's analysis of chimpanzee social structure (Wrangham 1975, 1979) indicated that the ultimate cause of that structure was ecological i.e., patchy food distribution leads to wide female dispersal for optimal foraging efficiency, which in turn favors a male kin breeding group that can maintain a territority that includes several individual female ranges. In contrast, Wrangham's phylogenetic model of the social structure of the stem-hominid (Wrangham, 1986) suggested that phylogenetic inertia may be partially responsible for the shared social features found among African Hominoidea. However, in the same work, Wrangham also suggested that further socioecological analysis of African apes may indicate whether food distribution and its effects on female dispersion/association may partially explain conservative African ape social features.Ghiglieri's phylogenetic model of the stem-hominid (1987, 1989), on the other hand, explained the conservative social features of bonobos, common chimpanzees, and hominids to be primarily a product of phylogenetic inertia and sexual selection. Furthermore, for Ghiglieri the most important sexual selection variable was a male communal reproductive strategy. This, according to Ghiglieri, is the ultimate cause of social structure. Notably, Ghiglieri (1984, 1985) had earlier stressed the overiding importance of a male communal reproductive strategy but was less dogmatic in his insistence that chimpanzees had essentially solved their ecological problems (e.g. that they had solved the food distribution problem by fusion-fission sociality; predators were never a real problem). Nevertheless, Ghiglieri's earlier position similarily expressed the idea that a communal reproductive strategy constituted the ultimate cause of social structure.The fourth objective was accomplished by presentation of an alternative model of chimpanzee social behavior which suggested that structure; the effect of phylogenetic inertia on social structure; chimpanzee social structure is the combined product of ecological and sexual selection forces: female optimal foraging, male mating strategies, and predator pressure. The model was considered by the author to be unique in that it integrated essential aspects of both Wrangham's and Ghiglieri's models and, in addition, provided support for Alexander's (1974) contention that predation pressure is an ultimate cause of ape social structure. The model also outlined scenarios for the evolution of chimpanzee group._ extensibility (fusion-fission sociality) and the capacity for warfare among chimpanzees.The last objective was achieved by a discussion of the implications that the author's model had for phylogenetic models of stem-hominid social structure. In this discussion the author reviewed the following issues as they related to the phylogenetic reconstruction of hominid social structure: the role of phylogeny and/or ecology in the causation of social encountered when using a phylogenetic referential model for the personal biases that enter into phylogenetic econstructions; pitfalls reconstruction of early hominid social evolution; the significance of chimpanzee models of social structure.The importance of the preceding study lay in its ability to stimulate improved conceptual models of African hominoid social structure. / Department of Anthropology

Grooming relationships between adult male chimpanzees are often reciprocal, i.e. individuals receive grooming from those they groom. Grooming may be reciprocated at the same time it is received (mutual grooming), or later within the same grooming session. Alternatively, it can be reciprocated at a much later stage, in another session. An analysis of individual grooming sessions at the dyadic level was used to investigate how chimpanzees reciprocate grooming within these sessions. This study describes the grooming and reciprocation of grooming by male chimpanzees, living in a multi-male, multi-female group at the Edinburgh Zoo, Scotland. A method for the analysis of dyadic grooming relationships was based on the presence or absence of mutual and unilateral grooming in a session, which allows seven types of grooming session to be distinguished. Grooming session was defined empirically, and the duration of the bout criterion interval (BCl) depended on the presence or absence of oestrous females. For comparison, however, the same BCI was used throughout. Without oestrous females, grooming was primarily reciprocated in sessions with mutual grooming and unilateral grooming by both participants. This kind of session proved highly cooperative and each male adjusted the duration of his unilateral grooming to that of mutual grooming, rather than to the duration of unilateral grooming by the other male. Mutual grooming was less important to dyads which had a strong grooming relationship. It is suggested that mutual grooming serves as an indication of the motivation to groom unilaterally. There was no indication that males reciprocated on the basis of TIT-FOR-TAT within these sessions, or between sessions in general. Alternative hypotheses of mutual grooming were only partly confirmed in that some dyads used mutual grooming to reduce the (already very short) time they spent in grooming. However, mutual grooming did not arise from the accidental overlap in the grooming of two partners. In the presence of oestrous females, grooming cooperation between the males broke down, and this was the result of heightened aggression as well as the presence of oestrous females itself. The balance in grooming given and received shifted in the direction of dominants (i.e. dominants received more) under the influence of oestrous females, but in the opposite direction under the influence of aggression. Feeding had no effect on the reciprocity of groormng. There was considerable dyadic variation. Some dyads groomed more when there were oestrous females, others groomed less. Some dyads had proportionally less mutual grooming with increasing numbers of oestrous females, others had more. There were generally no clear patterns of grooming reciprocation over longer time-spans than the session, but the overall degree of reciprocity of a dyad was frequently reached at the end of each day. Tracing the degree of reciprocation over a few weeks indicated that some dyads' grooming was governed by dominance, whereas that of others by cooperation.

Increasing and unsustainable demands on Africa's natural resources are having a profound effect on wild primate populations. Whilst wild populations are decreasing, numbers of orphaned primates, sanctuaries and attempts to reintroduce primates back to the natural environment, are increasing. Data were collected on the present status of African ape sanctuaries from questionnaires distributed to sanctuary managers. Across Africa there are 18 sanctuaries housing over 500 African great apes. Facilities and ideologies vary but the majority of sanctuaries profess a commitment to conservation through education, local capacity building, facilitating the enforcement of wildlife laws and other activities. From 1996 to 2001 the non-governmental organisation Habitat Ecologique et Liberte des Primates has released 37 wild-born chimpanzees(Pan troglodytes troglodytes) from an island sanctuary to mainland forest in the Conkouati-Douli National Park, Republic of Congo. Twenty-seven chimpanzees have been successfully reintroduced, three are known to have died and the status of seven remains unknown. This thesis investigated the behavioural adaptation of 15 of these released chimpanzees and reviews the reintroduction process employed. Analyses of post-release behavioural data revealed that activity budgets and diet were comparable to those of wild chimpanzees, and that seasonal variation influenced feeding behaviour and plant speciess election. The chimpanzees utilised both terrestrial and arboreal zones and all nested in trees. A number of recommendations are made for future reintroduction projects. These include selecting a release site that has no, or a low density of, wild conspecifics; developing a relationship of trust between chimpanzee and caretakers without excessive dependency; using the release site for pre-release training; use of radio telemetry; post-release support and monitoring. This study has revealed the many complex factors that are involved in the reintroduction process. Future attempts to reintroduce chimpanzees should be guided by the experiences and recommendations of the present study to maximise success.