Intracommunity Coalitionary Killing of an Adult Male Chimpanzee at Ngogo,Kibale National Park,Uganda

Intercommunity coalitionary killing of adult and adolescent males has been documented in two chimpanzee communities in the wild, and it was strongly suspected in a third. It may increase survivorship for the attackers, their mates, and their offspring by reducing the combined strength of hostile neighbors and/or by increasing territory size and food availability, and it may help the attackers to attract mates. Lethal coalitionary attacks by males on other male members of their own communities would not provide these benefits and are not expected, given the importance of cooperation among male community members in contests for dominance rank and in both defense and offense against neighboring males. Nevertheless, intracommunity coalitionary killings associated with struggles for alpha rank occur in the wild and in captivity, and observers have seen serious gang attacks on maturing adolescent and young adult males at Mahale and Budongo: the victim in the Budongo case was killed (Fawcett and Muhumuza, 2000). I describe a lethal attack on a young adult male by a large coalition of males from his community at Ngogo, Kibale National Park, Uganda. The Ngogo community is the largest known for chimpanzees and has an unusually large number of males. The attack was not related to a struggle for alpha rank: the victim was low-ranking and the community had a well-established alpha at the time. However, the victim had risen substantially in the male hierarchy over the past few years and might have appeared threatening to many higher-ranking males. Simultaneously, he associated relatively little with most other adult males, had relatively few grooming partners and was not well integrated into the male grooming network, and had no influential allies. The combination of these social factors with the unusual demographic circumstances – which presumably meant that mating competition was relatively high and the cost of losing one male relatively low – might have triggered the attack.     

New cases of inter-community infanticide by male chimpanzees at Ngogo,Kibale National Park,Uganda

Infanticide by males has been recorded in four chimpanzee populations, including that in Kibale National Park, Uganda. Some infanticidal attacks occur during inter-community aggression. The sexual selection hypothesis does not easily explain these attacks because they may not directly increase male mating opportunities. However, females in the attackers’ community may benefit by expanding their foraging ranges and thereby improving their reproductive success; thus infanticide may increase male reproductive success indirectly. We report two new cases of infanticide by male chimpanzees at Ngogo, Kibale National Park. Like two previous cases, these occurred during a boundary patrol and were almost certainly between-community infanticides. The patrolling males attacked despite the proximity of males from the victims’ presumed community. This probably explains why, unlike the earlier cases, they did not completely cannibalize their victims. Such attacks seem to be relatively common at Ngogo and infanticide may be an important source of infant mortality in neighboring communities. Our observations cannot resolve questions about the sexual selection hypothesis. However, they are consistent with the range expansion hypothesis: the infanticides occurred during a period of frequent encounters between communities associated with a mast fruiting event, and Ngogo community members greatly increased their use of areas near the attack site during another mast fruiting event one year later. Our observations contribute to growing evidence that lethal intergroup aggression is a common characteristic of wild chimpanzee populations.     

Violent coalitionary attacks and intraspecific killing in wild white-faced capuchin monkeys (<Emphasis Type="Italic">Cebus capucinus</Emphasis>)

During 12 years of observation, we have observed three confirmed and two inferred lethal coalitionary attacks on adult male white-faced capuchins (Cebus capucinus) by members of two habituated social groups at Lomas Barbudal Biological Reserve, Costa Rica. In one case, an alpha male was badly wounded and evicted from his group, and when later found by his former groupmates he was attacked by several of them and died less than 24 h later. In two other cases, lone extra-group males were mobbed by adult and immature males of a bisexual group. One victim's abdomen was torn open and he died less than 24 h later. A second victim was quite badly bitten but may have escaped. The fourth and fifth cases resulted from intergroup encounters. One victim lost the use of both arms but may have survived, whereas the other died of unknown causes within an hour of the attack. The observed death rate from coalitionary aggression at our site is approximately the same as that reported for eastern chimpanzees. Because at least three of the five observed incidents involved large coalitions attacking lone victims, they support the general hypothesis that imbalances of power contribute to intraspecific killing in primates. However, the occurrence of lethal coalitional attacks in a species lacking fission–fusion social organization poses a challenge to the more specific version of the imbalance-of-power hypothesis proposed by Manson and Wrangham in 1991 to explain chimpanzee and human intergroup aggression.     

Intra‐community coalitionary lethal attack of an adult male southern muriqui (Brachyteles arachnoides)

We report on the first evidence of intra‐community coalitionary lethal aggression in muriquis (Brachyteles). The event occurred in southern muriquis (Brachyteles arachnoides) during a long‐term study (>15 years) of two social groups inhabiting mostly pristine Atlantic forest habitat in the Parque Estadual Carlos Botelho, southern São Paulo State, Brazil. The attack took place deep in the core area of the Group Caetê home range. Tense agonistic behaviors and vocalizations preceded the lethal coalitionary attack, and the tension increased over a 36–48 hr period. One adult female and two unidentified individuals also took part in a coalition led by six adult males. The members of the coalition collectively approached, embraced, immobilized and repeatedly bit the entire body of an adult male, resulting in severe bleeding injuries and the victim's death in less than 1 hr after the attack commenced. Combined ecological, behavioral and spatial data related to the event indicate that this was an intra‐community attack and suggest social tensions related to mating competition as the proximate trigger of the coalitionary killing. The attack resembled those reported for chimpanzees, with clear numeric superiority and a low risk of injury to aggressors, resulting in the death of a lone conspecific victim. This observation (n=1) is suggestive of a capacity for escalated aggression in muriquis and reinforces arguments for the potential adaptive significance of intra‐community aggression in male philopatric societies, as reported for spider monkeys and chimpanzees. These characteristics challenge the view of the muriquis as a peaceful primate and support the general hypothesis that imbalances of power contribute to intra‐specific killing in primates, such as chimpanzees and humans. Am. J. Primatol. 71:860–867, 2009. © 2009 Wiley‐Liss, Inc.     

New Intercommunity Infanticides by the Chimpanzees of Ngogo,Kibale National Park,Uganda

Hostile intercommunity relations, including attacking and killing extra-community infants of both sexes have occurred at most wild chimpanzee sites. We describe three recent cases of intercommunity attacks on infants committed by members of the Ngogo chimpanzee community in Kibale National Park, Uganda. Two of the attacks resulted in confirmed infanticides while a third attack probably resulted in the infant's death. In common with previous accounts of chimpanzee infanticides, the attacks described here occurred during boundary patrols outside the Ngogo community's usual range, adult and adolescent males were the main participants, one infant was cannibalized after being killed, and the victims’ mothers did not accompany the attacking party back to the Ngogo range. However, the patrol parties during each infanticide were larger than before and included females from the Ngogo community. Our observations indirectly support both the range expansion and imbalance of power hypotheses, which address why and under which conditions chimpanzee intercommunity encounters lead to aggression. These cases of intercommunity infanticide add to the growing database of the phenomenon in wild chimpanzees.     

Demographic influences on the behavior of chimpanzees

Recent research has revealed substantial diversity in the behavior of wild chimpanzees. Understanding the sources of this variation has become a central focus of investigation. While genetic, ecological, and cultural factors are often invoked to explain behavioral variation in chimpanzees, the demographic context is sometimes overlooked as a contributing factor. Observations of chimpanzees at Ngogo, Kibale National Park, Uganda, reveal that the size and structure of the unit group or community can both facilitate and constrain the manifestation of behavior. With approximately 150 individuals, the Ngogo community is much larger than others that have been studied in the wild. We have taken advantage of the unusual demographic structure of this community to document new and intriguing patterns of chimpanzee behavior with respect to hunting, territoriality, and male social relationships. Chimpanzees at Ngogo hunt often and with a considerable degree of success. In addition, male chimpanzees there frequently patrol the boundary of their territory and engage in repeated bouts of lethal intergroup aggression. By forming two distinct subgroups, male chimpanzees at Ngogo also develop social bonds above the level of dyadic pairs. While the sheer number of chimpanzees contributes to differences in hunting, patrolling, mating, and subgrouping at Ngogo, the demographic situation may also constrain behavioral interactions. At Ngogo, male chimpanzees who are closely related genetically through the maternal line do not appear to affiliate or cooperate with each other. Demographic constraints may be responsible for this finding. In this paper, I use these examples to illustrate how the demographic context affects the possible range of behavioral options open to individuals and ultimately contributes to the explanation of behavioral diversity in chimpanzees.     

Fatal Chimpanzee Attack in Loango National Park,Gabon

In some populations, chimpanzees engage in lethal aggression within and between social units. We report a fatal attack on an adult male chimpanzee at a new research site in Loango National Park, Gabon. We found a fresh corpse of an adult male chimpanzee only a few hundred meters from the research camp, after noting numerous vocalizations and chimpanzee movements the previous evening. Previous contacts with chimpanzees and fresh tracks in the area around the corpse suggest that 2 communities of chimpanzees range where the attack occurred and that members of the neighboring community killed the chimpanzee. To support the conclusion, we conducted genetic analysis for 13 Y-chromosome loci and 9 microsatellite loci of fecal samples from the dead individual, 5 possible attackers, and 2 members of the other community Though we cannot exclude the possibility of an intracommunity killing, the combined observational and genetic evidence suggest an intercommunity attack. The case study adds to the growing evidence that intercommunity killings are a rare but widespread phenomenon among chimpanzees and not an artifact of human provisioning or habituation.     

Intergroup conflicts among chimpanzees in Taï National Park: lethal violence and the female perspective

Intercommunity aggression in chimpanzees and primitive warfare in humans possess striking similarities, such as the common occurrence of large male coalitions, systematic control of territory boundaries, and lethal attacks on isolated individuals from neighboring groups. However, an important apparent contrast is the absence of recurrent peaceful interactions between neighboring groups of chimpanzees. We observed a remarkable range of behavior in intergroup encounters among three habituated communities of chimpanzees in Taï National Park, Côte d'Ivoire. Lethal attacks are documented in these study groups for the first time, as well as year‐long exchanges of parous adult females and peaceful intergroup visits of mothers with infants. Demographic factors, including group size and number of adult males, are shown to affect the nature of intergroup interactions in ways not considered previously. A reconsideration of the difference in intergroup interactions between eastern and western chimpanzees is proposed including a more important consideration of the female's perspective. The inclusion of the new complexities in intergroup interactions in chimpanzees allows new parallels to be drawn with the evolution of primitive warfare in humans. Am. J. Primatol. 70:519–532, 2008. © 2008 Wiley‐Liss, Inc.     

Death of a wild chimpanzee community member: possible outcome of intense sexual competition

Our observations strongly support the view that a lethal gang attack occurred against a young adult male within a wild chimpanzee (Pan troglodytes schweinfurthii) community. This is the first report of such an occurrence. At least three adult male chimpanzees are believed to have been involved in the attack. This case is unusual because the attack was fatal, involved directed group aggression within a community, and was not linked to the overthrow of the alpha male. During the period before the attack, the percentage of cycling females in the community was very low (15 males:17 females, a maximum of two regularly cycling). This fact, coupled with the presence of one female in full genital tumescence, may have incited and escalated the violence of the attack. The lethal attack is interpreted as an act of intra-community male sexual competition resulting in the complete exclusion of one male from estrous females.     

New Cases of Intergroup Violence Among Chimpanzees in Gombe National Park,Tanzania

Despite considerable attention to chimpanzee intergroup violence, the number of observed cases remains small. We report 4 cases of intergroup violence that occurred in Gombe National Park, Tanzania, between 1993 and 2002. We observed (3 cases) or inferred (1 case) males from the Kasekela community to attack members of their 2 neighboring communities: Mitumba and Kalande. In 1993, Kasekela males killed and ate a female infant from Mitumba. In 1998, Kasekela males captured 2 infants (sex unknown) from Kalande, one of which escaped and the other was killed and eaten. Also in 1998, Kasekela males attacked an adolescent male from Kalande. The victim was alive but severely injured by the end of the attack. The intensity and duration of the attack are comparable to other attacks that resulted in fatal injuries. In 2002, observers found the body of an adolescent male from Mitumba following an incursion by Kasekela males into the area. The injuries inflicted on the Mitumba male together with circumstantial evidence suggest that Kasekela males killed him. The attacks support the view that intergroup violence is a persistent feature of chimpanzee societies and that the primary benefit attackers gain from them is reduced competition for resources.     

Infanticide and cannibalism by male chimpanzees at Ngogo, Kibale National Park, Uganda

Researchers have documented infanticide by adult males in four wild chimpanzee populations. Males in three of these have killed infants from outside of their own communities, but most infanticides, including one from Kanyawara, in Kibale National Park, Uganda, took place within communities. Here we report two new cases of infanticide by male chimpanzees at a second Kibale site, Ngogo, where the recently habituated chimpanzee community is the largest yet known. Both infanticides happended during boundary patrols, which occur at a high frequency there. Patrolling males attacked solitary females who were unable to defend their infants successfully. The victims were almost certainly not members of the Ngogo community. Males cannibalized both infants and completely consumed their carcasses. These observations show that infanticide by males is widespread in the Kibale population and that between-community infanticide also happens there. We discuss our observations in the context of the sexual selection hypothesis and other proposed explanations for infanticide by male chimpanzees. The observations support the arguments that infanticide has been an important selective force in chimpanzee social evolution and that females with dependent infants can be at great risk near range boundaries, but why male chimpanzees kill infants is still uncertain.     

Causes of death in the Kasekela chimpanzees of Gombe National Park, Tanzania

Understanding the rates and causes of mortality in wild chimpanzee populations has important implications for a variety of fields, including wildlife conservation and human evolution. Because chimpanzees are long-lived, accurate mortality data requires very long-term studies. Here, we analyze 47 years of data on the Kasekela community in Gombe National Park. Community size fluctuated between 38 and 60, containing 60 individuals in 2006. From records on 220 chimpanzees and 130 deaths, we found that the most important cause of mortality in the Kasekela community was illness (58% of deaths with known cause), followed by intraspecific aggression (20% of deaths with known cause). Previous studies at other sites also found that illness was the primary cause of mortality and that some epidemic disease could be traced to humans. As at other study sites, most deaths due to illness occurred during epidemics, and the most common category of disease was respiratory. Intraspecific lethal aggression occurred within the community, including the killing of infants by both males and females, and among adult males during the course of dominance-related aggression. Aggression between communities resulted in the deaths of at least five adult males and two adult females in the Kasekela and Kahama communities. The frequency of intercommunity violence appears to vary considerably among sites and over time. Intercommunity lethal aggression involving the Kasekela community was observed most frequently during two periods. Other less common causes of death included injury, loss of mother, maternal disability, and poaching.     

Why do chimpanzee males attack the females of neighboring communities?

Our closest nonhuman primate relatives, chimpanzees, engage in potentially lethal between‐group conflict; this collective aggressive behavior shows parallels with human warfare. In some communities, chimpanzee males also severely attack and even kill females of the neighboring groups. This is surprising given their system of resource defense polygyny, where males are expected to acquire potential mates. We develop a simple mathematical model based on reproductive skew among primate males to solve this puzzle. The model predicts that it is advantageous for high‐ranking males but not for low‐ranking males to attack females. It also predicts that more males gain a benefit from attacking females as the community's reproductive skew decreases, i.e., as mating success is more evenly distributed. Thus, fatal attacks on females should be concentrated in communities with low reproductive skew. These attacks should also concur with between‐community infanticide. A review of the chimpanzee literature provides enough preliminary support for this prediction to warrant more detailed testing. Am J Phys Anthropol 155:430–435, 2014. © 2014 Wiley Periodicals, Inc.     

Grooming Between Male Chimpanzees at Ngogo,Kibale National Park. I. Partner Number and Diversity and Grooming Reciprocity

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.     

The development of affiliative and coercive reproductive tactics in male chimpanzees

Like many animals, adult male chimpanzees often compete for a limited number of mates. They fight other males as they strive for status that confers reproductive benefits and use aggression to coerce females to mate with them. Nevertheless, small-bodied, socially immature adolescent male chimpanzees, who cannot compete with older males for status nor intimidate females, father offspring. We investigated how they do so through a study of adolescent and young adult males at Ngogo in Kibale National Park, Uganda. Adolescent males mated with nulliparous females and reproduced primarily with these first-time mothers, who are not preferred as mating partners by older males. Two other factors, affiliation and aggression, also influenced mating success. Specifically, the strength of affiliative bonds that males formed with females and the amount of aggression males directed toward females predicted male mating success. The effect of male aggression toward females on mating success increased as males aged, especially when they directed it toward females with whom they shared affiliative bonds. These results mirror sexual coercion in humans, which occurs most often between males and females involved in close, affiliative relationships.     

Hormonal Mechanisms for Regulation of Aggression in Human Coalitions

Coalitions and alliances are core aspects of human behavior. All societies recognize alliances among communities, usually based in part on kinship and marriage. Aggression between groups is ubiquitous, often deadly, fueled by revenge, and can have devastating effects on general human welfare. Given its significance, it is surprising how little we know about the neurobiological and hormonal mechanisms that underpin human coalitionary behavior. Here we first briefly review a model of human coalitionary behavior based on a process of runaway social selection. We then present several exploratory analyses of neuroendocrine responses to coalitionary social events in a rural Dominican community, with the objective of understanding differences between in-group and out-group competition in adult and adolescent males. Our analyses indicate: (1) adult and adolescent males do not elevate testosterone when they defeat their friends, but they do elevate testosterone when they defeat outsiders; (2) pre-competition testosterone and cortisol levels are negatively associated with strength of coalitionary ties; and (3) adult males usually elevate testosterone when interacting with adult women who are potential mates, but in a striking reversal, they have lower testosterone if the woman is a conjugal partner of a close friend. These naturalistic studies hint that reciprocity, dampening of aggression, and competition among friends and allies may be biologically embedded in unique ways among humans.     

Demographic influences on the hunting behavior of chimpanzees

We investigated hunting in an unusually large community of wild chimpanzees at Ngogo in the Kibale National Park, Uganda. Aspects of predation were recorded with respect to the prey, the predators, and hunting episodes. During 23 months of observation, the Ngogo chimpanzees caught 128 prey items from four primate and three ungulate species. Chimpanzees preyed selectively on immature red colobus primarily during group hunts, with adult males making the majority of kills. Party size and composition were significant predictors of the probability that chimpanzees would hunt and of their success during attempts. Chimpanzees were more likely to hunt red colobus if party size and the number of male hunters were large; party size and the number of male hunters were also significantly larger in successful compared with unsuccessful hunts. The Ngogo chimpanzees did not appear to hunt cooperatively, but reciprocal meat-sharing typically took place after kills. Hunts occurred throughout the year, though there was some seasonality as displayed by periodic hunting binges. The extremely high success rate and large number of kills made per successful hunt are the two most striking aspects of predation by the Ngogo chimpanzees. We compare currently available observations of chimpanzee hunting behavior across study sites and conclude that the large size of the Ngogo community contributes to their extraordinary hunting success. Demographic differences between groups are likely to contribute to other patterns of interpopulation variation in chimpanzee predation. Am J Phys Anthropol 109:439–454, 1999. © 1999 Wiley-Liss, Inc.     

Lethal intragroup aggression by adult male spider monkeys (Ateles geoffroyi)

I report three cases of coalitionary aggression by adult male black-handed spider monkeys (Ateles geoffroyi) against subadult males within their community on Barro Colorado Island (BCI), Panama. Two of these cases were followed by the disappearance and presumed death of the victim. Similar behavior was recently reported by Valero et al. [in press], who suggested that this behavior may be the result of intense male reproductive competition. Like the single instance they reported, the cases I report all occurred when the operational sex ratio was approximately 1:1, which suggests that intense competition among males for access to reproductively viable females may be a contributing factor. Additionally the very low density of spider monkeys on BCI may play a significant role in the occurrence of this lethal aggression. Large numbers of adult males are not necessary to protect a territorial boundary against neighboring groups, and additional males may act merely as mating competition.     

Intelligence, Coalitional Killing, and the Antecedents of War

Advances in primatological research have recently led to a hypothesis that lethal coalitionary raiding in chimpanzees is the product of an evolutionarily adaptive "dominance drive" that disposes adult males to seek out low-cost opportunities for conspecific killing. This conclusion has been extended into a claim that human warfare and other forms of coalitional killing are outcomes of a hardwired, "demonic male" complex. Reversing this evidential approach, I argue from data on conspecific killing in humans that humans and chimpanzees have an aversion to killing conspecifics. Their lethal violence, I propose, is more parsimoniously explained as the result of a developed intelligence capable of envisioning the future and, when necessary, of disabling this aversion to achieve desired goals.     

Intergroup Aggression in Chimpanzees and War in Nomadic Hunter-Gatherers

Chimpanzee and hunter-gatherer intergroup aggression differ in important ways, including humans having the ability to form peaceful relationships and alliances among groups. This paper nevertheless evaluates the hypothesis that intergroup aggression evolved according to the same functional principles in the two species—selection favoring a tendency to kill members of neighboring groups when killing could be carried out safely. According to this idea chimpanzees and humans are equally risk-averse when fighting. When self-sacrificial war practices are found in humans, therefore, they result from cultural systems of reward, punishment, and coercion rather than evolved adaptations to greater risk-taking. To test this “chimpanzee model,” we review intergroup fighting in chimpanzees and nomadic hunter-gatherers living with other nomadic hunter-gatherers as neighbors. Whether humans have evolved specific psychological adaptations for war is unknown, but current evidence suggests that the chimpanzee model is an appropriate starting point for analyzing the biological and cultural evolution of warfare.