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How can collective action evolve when individuals benefit from cooperation regardless of whether they pay its participation costs? According to one influential perspective, collective action problems are common, especially when groups are large, but may be solved when individuals who have more to gain from the collective good or can produce it at low costs provide it to others as a byproduct. Several results from a 20-y study of one of the most striking examples of collective action in nonhuman animals, territorial boundary patrolling by male chimpanzees, are consistent with these ideas. Individuals were more likely to patrol when (i) they had more to gain because they had many offspring in the group; (ii) they incurred relatively low costs because of their high dominance rank and superior physical condition; and (iii) the group size was relatively small. However, several other findings were better explained by group augmentation theory, which proposes that individuals should bear the short-term costs of collective action even when they have little to gain immediately if such action leads to increases in group size and long-term increases in reproductive success. In support of this theory, (i) individual patrolling effort was higher and less variable than participation in intergroup aggression in other primate species; (ii) males often patrolled when they had no offspring or maternal relatives in the group; and (iii) the aggregate patrolling effort of the group did not decrease with group size. We propose that group augmentation theory deserves more consideration in research on collective action. [ABSTRACT FROM AUTHOR]

Male chimpanzees, Pan troglodytes, are well known for affiliating and cooperating in a variety of behavioural contexts. Prior field research indicates that maternal kinship does not affect patterns of affiliation and cooperation by males in the same social group. Two questions remain unclear from this finding. First, why do male chimpanzees not bias their behaviour towards maternal kin? Second, what factors account for who affiliates and cooperates with whom? We conducted behavioural observations of an unusually large community of chimpanzees at Ngogo, Kibale National Park, Uganda, to test the hypothesis that demographic constraints limit the number of maternal kin with whom male chimpanzees can cooperate, and thereby lead them to form selective bonds with nonkin of similar age and status. Results indicated that male age and rank are significantly associated with four measures of social behaviour. Members of the same age class and individuals close in rank were more likely to affiliate and cooperate than males that belonged to different age and rank classes. Additional analyses replicate earlier findings and show that males who affiliated and cooperated were not closely related through the maternal line, as assayed by mtDNA haplotype sharing. These results add to our growing understanding of the important role demographic and social constraints play in animal behaviour. [Copyright &y& Elsevier]

Chimpanzees (Pan troglodytes ) prey on a variety of vertebrates, mostly on red colobus (Procolobus spp. ) where the two species are sympatric. Variation across population occurs in hunting frequency and success, in whether hunting is cooperative, i.e., payoffs to individual hunters increase with group size, and in the extent to which hunters coordinate their actions in space and time, and in the impact of hunting on red colobus populations. Also, hunting frequency varies over time within populations, for reasons that are unclear. We present new data on hunting by chimpanzees at Ngogo, Kibale National Park, Uganda, and combine them with earlier data (Mitani and Watts, 1999, Am. J. Phys. Anthropol. 109: 439–454) to examine hunting frequency and success, seasonality, and cooperation. The Ngogo community is the largest and has the most males of any known community. Chimpanzees there mostly hunt red colobus and are much more successful and make many more kills per hunt than at other sites; they kill 6–12% of the red colobus population annually. The number of kills and the offtake of meat per hunt increase with the number of hunters, but per capita meat intake is independent of hunting party size; this suggests that cheating occurs in large parties. Some behavioral cooperation occurs. Hunting success and estimated meat intake vary greatly among males, partly due to dominance rank effects. The high overall success rate leads to relatively high average per capita meat intake despite the large number of consumers. The frequency of hunts and of hunting patrols varies positively with the availability of ripe fruit; this is the first quantitative demonstration of a relationship between hunting frequency and the availability of other food, and implies that the chimpanzees hunt most when they can easily meet energy needs from other sources. We provide the first quantitative support for the argument that variation in canopy structure influences decisions to hunt red colobus because hunts are easier where the canopy is broken. [ABSTRACT FROM AUTHOR]

Investigates the factors that influence chimpanzee hunting decisions and meat sharing. Compensation for seasonal shortages in food availability; Use of meat as a social tool to develop and maintain alliances with other males; Use of meat in exchange for matings; Frequency of hunting.

Chimpanzees are among the few mammals that engage in lethal coalitionary aggression between groups. Most attacks on neighbors occur when parties made up mostly of adult males patrol boundaries of their community's range. Patrols have time, energy, and opportunity costs, and entail some risks despite the tendency of males to attack only when they greatly outnumber their targets. These factors may lead to a collective action problem. Potential benefits include protection of community members, particularly infants; range expansion and increases in the amount and quality of food available; and incorporation of more females into the community. Males may not share these equally; for example, those able to obtain large shares of matings may stand to gain most by participating in patrols and to lose most by refraining. Despite the attention that boundary patrolling has attracted, few relevant quantitative data are available. Here, we present detailed data on boundary patrolling and intergroup aggression in a chimpanzee community at Ngogo, Kibale National Park, Uganda, that is unusually large and has more males than any other known community. Males there patrolled much more often, and patrol parties were much larger on average, than at two other sites for which comparative data exist. Our findings support the argument that male participation varies along with variation in potential gains, in willingness to take risks, and in skill at handling these risks. Both the overall frequency with which individual males patrolled and their willingness to join patrols as others set off on them were positively associated with variation in mating success, in participation in hunts of red colobus monkeys, and in hunting success. Males patrolled relatively often with others with whom they associated often in general, with whom they often groomed, and with whom they formed coalitions in withincommunity agonism. This indicates that they were most willing to take risks associated with patrolling when with others they trusted to take the same risks. [ABSTRACT FROM AUTHOR]

Allogrooming contributes to the development and maintenance of social relationships, including those that involve alliances, in many primate species. Variation in relatedness, dominance rank, and other factors can produce variation in the value of others as grooming partners. Several models have been developed to account for variation in the distribution of grooming in relation to dominance ranks. These start from the premise that individuals are attracted to high-ranking partners, but time limits, direct competition, and prior grooming engagement between high-ranking individuals can constrain access to them. Sambrook et al. (1995) formalized some of these models and showed the importance of taking group size variation into account when assessing them. Chimpanzees form multimale communities in which males are the philopatric sex. Males commonly associate and groom with each other; they also form dominance hierarchies and form alliances that influence dominance ranks and mating success. Both male rank and the rank distance between partners are significantly correlated with the distribution of grooming between males in an extremely large chimpanzee community at Ngogo, Kibale National Park, Uganda, that has more males than any other known community. High-ranking males had more grooming partners than mid- or low-ranking males. Grooming predominantly went up the dominance hierarchy, but was also concentrated among males that were close in rank. Rank and rank distance apparently both affected grooming independently of reciprocity in grooming and independently of the frequency with which males associated in temporary parties. However, the data do not clearly indicate how constraints on access to partners might have operated. Published data from a smaller chimpanzee community at Mahale show no rank or rank distance effect on male grooming. These results and earlier, conflicting findings on the association between dominance rank and grooming in male chimpanzees indicate that variation in group size, i.e., the number of males per community, probably influences the strength of any such effects, as happens for grooming between females in several cercopithecine species. Data on coalitions at Ngogo support the argument that high-ranking males are valuable social partners, and similarity in strategies of alliance formation may influence the distribution of grooming. [ABSTRACT FROM AUTHOR]

Allogrooming serves many social functions in primates. Grooming can help individuals to service social relationships generally, sometimes reciprocally, and may be particularly important in the development and maintenance of alliances. However, time constraints limit the number of partners with whom one individual can groom enough to maintain cooperative relationships. As a result, the size of its grooming network may reach an asymptote as the size of its group increases, and it may distribute its grooming less equally among potential partners. Chimpanzees live in multimale, fission-fusion communities; males are philopatric, and commonly associate and groom with each other. Males form within-community alliances that influence dominance rank and access to mates, and allies groom with each other regularly; males also cooperate in aggression between communities. The chimpanzee community at Ngogo, in Kibale National Park, Uganda, is unusually large and has more males than any other known community. Field data show that adult Ngogo males groomed far more with other adult males than with females or with adolescent males, in contrast to a previous report (Ghiglieri, 1984). Adolescent males groomed adults much more than the reverse; males groomed and were groomed by females about equally. Individual males groomed mostly with a small number of other males. On average, males at Ngogo had only slightly more male grooming partners overall and had the same number of important partners as those of males in a much smaller community in the Mahale National Park, Tanzania, and they distributed their grooming less equitably. These results fit those expected if limits on available grooming time cause males to have a loyalty problem as the number of potential grooming and alliance partners increases. Despite differences in the extent and equitability of their grooming networks, males at both Ngogo and Mahale showed reciprocity in grooming. Grooming reciprocity has been demonstrated for captive chimpanzee males, but the Ngogo findings are the first demonstrations of reciprocity in wild communities. [ABSTRACT FROM AUTHOR]

Presents information on a study which investigated the demographic effects on the hunting behavior of a community of wild chimpanzees at Ngogo in the Kibale National Park, Uganda. Methods; Results; Discussion and comparisons with other studies.

Cooperative mate guarding by males is unusual in mammals and birds, largely because fertilizations are non-shareable. Chimpanzees live in fission-fusion communities that have cores of philopatric males who cooperate in inter-group aggression and in defending access to the females in their community. Male contest mating competition is restrained within communities, but single high-ranking males sometimes try to mate guard estrous females. Data from an unusually large chimpanzee commmunity at Ngogo, Kibale National Park, Uganda, that contains more males than any previously studied community show new variation in chimpanzee mate-guarding behavior. Contrary to expectation given the large number of males, mate guarding was as common as, or more common than, at other sites, and males other than the alpha male guarded more often. More strikingly, pairs or trios of top-ranking males sometimes engaged in cooperative aggression to prevent estrous females from mating with other males, but tolerated each other's mating activities. Both single males and coalitions mostly guarded periovulatory females. Mate-guarding coalitions were previously unknown in chimpanzees. Coalitions occurred in large mating parties, seemingly because these often contained too many males for single males to maintain exclusive access to estrous females. Coalition members gained higher shares of copulations than they could have expected from solo mate guarding, and suffered lower per capita costs of guarding (as inferred from aggression rates). Two males who most often participated in coalitions formed two-male coalitions at about the point where the number of males present made it unlikely that either could get 50% or more of total copulations on his own, and formed trios when this value dropped below 33%. Kin selection could be a factor in cooperation among male chimpanzees, but coalition members were not necessarily close relatives and the apparent structure of payoffs fit that of mutualism. Furthermore, reliance of male chimpanzees on support from allies to maintain high rank could have led to trading of mating exclusivity for support against mating competitors. [ABSTRACT FROM AUTHOR]

Summary: Chimpanzees make lethal coalitionary attacks on members of other groups . This behavior generates considerable attention because it resembles lethal intergroup raiding in humans . Similarities are nevertheless difficult to evaluate because the function of lethal intergroup aggression by chimpanzees remains unclear. One prominent hypothesis suggests that chimpanzees attack neighbors to expand their territories and to gain access to more food . Two cases apparently support this hypothesis, but neither furnishes definitive evidence. Chimpanzees in the Kasekela community at Gombe National Park took over the territory of the neighboring Kahama community after a series of lethal attacks . Understanding these events is complicated because the Kahama community had recently formed by fissioning from the Kasekela group and members of both communities had been provisioned with food. In a second example from the Mahale Mountains, the M group chimpanzees acquired part of the territory of the adjacent K group after all of the adult males in the latter disappeared . Although fatal attacks were suspected from observations of intergroup aggression, they were not witnessed, and as a consequence, this case also fails to furnish conclusive evidence. Here we present data collected over 10 years from an unusually large chimpanzee community at Ngogo, Kibale National Park, Uganda. During this time, we observed the Ngogo chimpanzees kill or fatally wound 18 individuals from other groups; we inferred three additional cases of lethal intergroup aggression based on circumstantial evidence (see ). Most victims were caught in the same region and likely belonged to the same neighboring group. A causal link between lethal intergroup aggression and territorial expansion can be made now that the Ngogo chimpanzees use the area once occupied by some of their victims. [Copyright &y& Elsevier]