increasing their access to key resources, such as food or mates.1-5 Alternatively, it has 5 been argued to be a non-adaptive result of human impacts, such as habitat destruction 6 or provisioning of food. [6][7][8][9] To discriminate between these hypotheses we compiled long-7 term information from 18 chimpanzee communities and 4 bonobo communities. Our 8 data include 152 killings (N=58 observed, 41 inferred, and 53 suspected killings) by 9 chimpanzees in 15 communities and one suspected killing by bonobos. We found that 10 males had the greatest involvement as attackers (92% of participants) and victims 11 (73%); most killings (66%) involved intercommunity attacks; and attackers greatly 12 outnumbered their victims (median 8:1 ratio). Variation in rates of killing among 13communities depended on demographic variables but was unrelated to measures of 14 human impacts. These results from all major study populations over the last five 15 decades are consistent with previously proposed adaptive explanations for killing by 16 chimpanzees but not with the human impact hypothesis. 17 18Conspecific killing has been documented at multiple chimpanzee study sites, 2-5,10-12 but rates 19 vary greatly among sites. The human impact hypothesis and the adaptive strategies 20 hypothesis yield contrasting predictions, which we test here (Tables 1, 2). The human impact 21 hypothesis states that killing occurs mainly as an incidental outcome of aggression, 22 exacerbated by human activities such as providing a concentrated food resource, 23 deforestation-induced crowding, anthropogenic diseases or hunting. Accordingly, lethal 24 aggression should be high where human disturbance is high. In contrast, the adaptive strategies hypothesis views aggression as an evolved strategic 27 response by which aggressors tend to increase their fitness through increased access to 28 territory, food, mates or other benefits. [1][2][3][4][5][10][11][12][13][14][15][16][17] 45Intracommunity infanticide by females may result from intense competition among females 46 for the best feeding areas.17 Population differences in rates of killing are accordingly 47 expected to result from socioecological factors such as differences in grouping patterns 2,11 48 and/or demography.14 Lethal aggression thus occurs within a diverse set of circumstances, 49 but is expected to be most commonly committed by males; directed towards males; directed 50 6 towards non-kin, particularly members of other groups; and committed when overwhelming 51 numerical superiority reduces the costs of killing. 52 53Previous studies have developed and tested these specific hypotheses 2,5,[11][12][13][14][15][16][17] ; the present study 54 represents the first effort to test multiple hypotheses simultaneously with a comprehensive 55 dataset. To do so, we assembled data from 18 chimpanzee communities from both eastern 56 (N=12) and western (N=6) clades 24 of chimpanzees studied over 426 years (median = 21 57 years; range: 4-53) and from 4 bonobo communities studied for 92 years (media...

The forests surrounding Bossou, Guinea, are home to a small, semi-isolated chimpanzee community studied for over three decades [1]. In 1992, Matsuzawa [2] reported the death of a 2.5-year-old chimpanzee (Jokro) at Bossou from a respiratory illness. The infant's mother (Jire) carried the corpse, mummified in the weeks following death, for at least 27 days. She exhibited extensive care of the body, grooming it regularly, sharing her day- and night-nests with it, and showing distress whenever they became separated. The carrying of infants' corpses has been reported from a number of primate species, both in captivity and the wild [3-7] - albeit usually lasting a few days only - suggesting a phylogenetic continuity for a behavior that is poignant testament to the close mother-infant bond which extends across different primate taxa. In this report we recount two further infant deaths at Bossou, observed over a decade after the original episode but with striking similarities.

The construction of nests (or beds) for sleeping is a chimpanzee universal, yet little is known about the adaptive function of nest-building. We present an indepth study of nest-building by unhabituated chimpanzees at the Seringbara study site in the Nimba Mountains, Guinea, West Africa. We recorded 1520 chimpanzee nests over 28 mo during three study periods between 2003 and 2008. We investigated where chimpanzees built their nests, both across the home range and in nest trees, and assessed how altitude and habitat type affected nest site selectivity. We examined whether or not chimpanzees were selective in nest tree choice regarding physical tree characteristics and tree species and assessed plant species preference for both treeand ground-nesting. We tested three, nonmutually exclusive, hypotheses for the function of arboreal nest-building. We assessed whether selectivity for nest tree characteristics reflected an antipredator strategy, examined whether nesting patterns (both arboreal and terrestrial) and nesting height were influenced by variation in climatic conditions (temperature, humidity, wind), and measured mosquito densities at ground level and in trees at 10 m and related mosquito densities to nesting patterns. Chimpanzees preferred to nest above 1000 m and nested mainly in primary forest. They preferred relatively large trees with a low first branch, dense canopy, and small leaves and showed preference for particular tree species, which was stable across years, whereas plant choice for ground-nesting was largely based on plant availability. We found no support for the antipredation hypothesis, nor did mosquito densities Int explain arboreal nest-building. The thermoregulation hypothesis was supported, as both nesting patterns and nest-height variation across seasons reflected a humidity-avoidance strategy. Chimpanzees nested higher in trees and at higher altitudes in the wet season. In sum, chimpanzees were selective in their choice of nest sites, locations, and materials, and tree-nesting patterns at Seringbara were best explained by a thermoregulation strategy of humidity avoidance.

Tool use in extant primates may inform our understanding of the conditions that favoured the expansion of hominin technology and material culture. The 'method of exclusion' has, arguably, confirmed the presence of culture in wild animal populations by excluding ecological and genetic explanations for geographical variation in behaviour. However, this method neglects ecological influences on culture, which, ironically, may be critical for understanding technology and thus material culture. We review all the current evidence for the role of ecology in shaping material culture in three habitual tool-using non-human primates: chimpanzees, orangutans and capuchin monkeys. We show that environmental opportunity, rather than necessity, is the main driver. We argue that a better understanding of primate technology requires explicit investigation of the role of ecological conditions. We propose a model in which three sets of factors, namely environment, sociality and cognition, influence invention, transmission and retention of material culture.

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The chimpanzees (Pan troglodytes verus) of the Nimba Mountains, Guinea, West Africa, commonly make both elaborate ("night") and simple ("day") nests on the ground. In this study we investigated which factors might influence ground-nesting in this population, and tested two ecological hypotheses: 1) climatic conditions, such as high wind speeds at high altitudes, may deter chimpanzees from nesting in trees; and 2) a lack of appropriate arboreal nesting opportunities may drive the chimpanzees to nest on the ground. In addition to testing these two hypotheses, we explored whether ground-nesting is a sex-linked behavior. Data were collected monthly between August 2003 and May 2004 along transects and ad libitum. To identify the sex of ground-nesting individuals, we used DNA extracted from hair samples. The results showed that the occurrence and distribution of ground nests were not affected by climatic conditions or a lack of appropriate nest trees. Support was found for the notion that ground-nesting is a sex-linked behavior, as males were responsible for building all of the elaborate ground nests and most of the simple ground nests sampled. Elaborate ground nests occurred mostly in nest groups associated with tree nests, whereas simple ground nests usually occurred without tree nests in their vicinity. These results suggest that ground-nesting may be socially, rather than ecologically, determined.

Reconciliation is a conflict resolution mechanism that is common to many gregarious species with individualized societies. Reconciliation repairs the damaged relationship between the opponents and decreases postconflict (PC) anxiety. The "integrated hypothesis" links the quality of the opponents' relationship to PC anxiety, since it proposes that conflicts among partners with high relationship quality will yield high levels of PC anxiety, which in turn will lead to an increased likelihood of reconciliation. We tested the integrated hypothesis in captive chimpanzees (Pan troglodytes) in the Arnhem Zoo, The Netherlands. We applied the standard PC/matched control (MC) method. Our results mostly support the integrated hypothesis, in that more valuable and compatible partners (i.e., males and frequent groomers) reconciled more often than less valuable and weakly compatible partners (i.e., females and infrequent groomers). In addition, PC anxiety was higher after conflicts among males than among females. Emotional arousal thus appears to be a mediator facilitating reconciliation. However, in contrast to the predictions derived from the integrated hypothesis, PC anxiety appeared only in aggressees, and not in aggressors, of conflicts. This suggests that while relationship quality determines PC anxiety, it is dependent on the role of the participants in the conflict.

One contribution of 15 to a theme issue 'Innovation in animals and humans: understanding the origins and development of novel and creative behaviour'. Young orangutans are highly neophobic, avoid independent exploration and show a preference for social learning. Accordingly, they acquire virtually all their learned skills through exploration that is socially induced. Adult exploration rates are also low. Comparisons strongly suggest that major innovations, i.e. behaviours that have originally been brought into the population through individual invention, are made where ecological opportunities to do so are propitious. Most populations nonetheless have large innovation repertoires, because innovations, once made, are retained well through social transmission. Wild orangutans are therefore not innovative. In striking contrast, zoo-living orangutans actively seek novelty and are highly exploratory and innovative, probably because of positive reinforcement, active encouragement by human role models, increased sociality and an expectation of safety. The explanation for this contrast most relevant to hominin evolution is that captive apes generally have a highly reduced cognitive load, in particular owing to the absence of predation risk, which strongly reduces the costs of exploration. If the orangutan results generalize to other great apes, this suggests that our ancestors could have become more curious once they had achieved near-immunity to predation on the eve of the explosive increase in creativity characterizing the Upper Palaeolithic Revolution.