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...

Domestic dogs (Canis familiaris) and great apes from the genus Pan were tested on a series of object choice tasks. In each task, the location of hidden food was indicated for subjects by some kind of communicative, behavioral, or physical cue. On the basis of differences in the ecologies of these 2 genera, as well as on previous research, the authors hypothesized that dogs should be especially skillful in using human communicative cues such as the pointing gesture, whereas apes should be especially skillful in using physical, causal cues such as food in a cup making noise when it is shaken. The overall pattern of performance by the 2 genera strongly supported this social-dog, causal-ape hypothesis. This result is discussed in terms of apes' adaptations for complex, extractive foraging and dogs' adaptations, during the domestication process, for cooperative communication with humans.

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Observations of hunting and meat eating in our closest living relatives, chimpanzees (Pan troglodytes), suggest that among primates, regular inclusion of meat in the diet is not a characteristic unique to Homo. Wild chimpanzees are known to consume vertebrate meat, but its actual dietary contribution is, depending on the study population, often either unknown or minimal. Constraints on continual direct observation throughout the entire hunting season mean that behavioral observations are limited in their ability to accurately quantify meat consumption. Here we present direct stable isotope evidence supporting behavioral observations of frequent meat eating among wild adult male chimpanzees (Pan troglodytes verus) in Taï National Park, Côte d'Ivoire. Meat eating among some of the male chimpanzees is significant enough to result in a marked isotope signal detectable on a short-term basis in their hair keratin and long-term in their bone collagen. Although both adult males and females and juveniles derive their dietary protein largely from daily fruit and seasonal nut consumption, our data indicate that some adult males also derive a large amount of dietary protein from hunted meat. Our results reinforce behavioral observations of male-dominated hunting and meat eating in adult Taï chimpanzees, suggesting that sex differences in food acquisition and consumption may have persisted throughout hominin evolution, rather than being a recent development in the human lineage.dietary ecology | stable isotope analysis | human evolution C omparisons with extant primates and other mammals are essential to understanding the varied ecological niches occupied by early hominins. Data from chimpanzees (Pan troglodytes) indicate that populations living in different forest habitats have different food repertoires (1-4). Variation in hunting behavior and meat consumption has been observed between populations, with chimpanzee communities acquiring and consuming meat with varying levels of importance, from those who hunt rarely and largely opportunistically for slow-moving small mammals (4) to more regular, systematic hunting of medium-sized prey (2, 3, 5). As all chimpanzee populations rely heavily on various plants, nuts, and invertebrates for their daily energy requirements, the disparity in vertebrate meat consumption across populations has led some researchers (6, 7) to suggest that meat is an occasionally consumed, nonessential dietary supplement.The chimpanzees (Pan troglodytes verus) of the Taï National Park, Côte d'Ivoire, are known to be some of the most specialized chimpanzee hunters, consuming large quantities of meat annually, predominantly from Western red colobus (Procolobus badius) and, occasionally, Western black-and-white colobus (Colobus polykomos) monkeys (3,8). Hunting at Taï is cooperative among the male chimpanzees, and after capture, division of the resultant prey rewards participation in the hunt, rather than nepotism (2, 8). A significant adult male sex bias is also evident, with adult males reported to consum...