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New Cases of Intergroup Violence Among Chimpanzees in Gombe ...

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544 Wilson, Wallauer, and Pusey<br />

range <strong>of</strong> the Mitumba community. In the observed cases, the attackers had<br />

overwhelm<strong>in</strong>g numerical superiority, with parties conta<strong>in</strong><strong>in</strong>g ≥4 adult males<br />

attack<strong>in</strong>g victims that were either alone or limited to mothers and <strong>in</strong>fants. The<br />

attackers suffered no more than superficial <strong>in</strong>juries <strong>in</strong> any case. The attackers<br />

spent little time feed<strong>in</strong>g <strong>in</strong> the border regions, suggest<strong>in</strong>g they traveled to<br />

border regions to search for neighbors rather than to feed. The features<br />

shared by the attacks are thus broadly consistent with previous descriptions<br />

<strong>of</strong> lethal raid<strong>in</strong>g <strong>in</strong> chimpanzees (Manson and Wrangham, 1991).<br />

Most or all males present, and to some extent the females, participated<br />

<strong>in</strong> they attacks, <strong>in</strong>dicat<strong>in</strong>g that they were not the outcome <strong>of</strong> <strong>in</strong>dividual<br />

idiosyncrasy or pathology. Nonetheless, males varied <strong>in</strong> the frequency with<br />

which they were present for attacks, with 3 specific males known or <strong>in</strong>ferred<br />

to be present <strong>in</strong> all 4 cases (Frodo, Pax and Wilkie; Table I). Another 6 males<br />

were present for ≥2 attacks (Apollo, Beethoven, Freud, Gimble, Gobl<strong>in</strong> and<br />

Tubi).<br />

Although the sample <strong>of</strong> observed <strong>in</strong>tergroup attacks rema<strong>in</strong>s sufficiently<br />

small and diverse to preclude def<strong>in</strong>itive hypothesis test<strong>in</strong>g, the observations<br />

help to clarify which <strong>of</strong> 3 proposed hypothesis—sexual selection,<br />

rival coalition reduction, and resource competition—best expla<strong>in</strong>s the benefits<br />

ga<strong>in</strong>ed by attackers.<br />

First, the sexual selection hypothesis predicts that, <strong>in</strong> the case <strong>of</strong> <strong>in</strong>tergroup<br />

<strong>in</strong>fanticide, the mothers <strong>of</strong> the <strong>in</strong>fanticide victims should be more<br />

likely to mate with the attackers <strong>in</strong> the future. Neither <strong>of</strong> the <strong>in</strong>fanticides<br />

was followed by observed mat<strong>in</strong>g between the mother and the killers <strong>of</strong> the<br />

<strong>in</strong>fant, but only <strong>in</strong> the case <strong>of</strong> the Mitumba female Rafiki was the mother<br />

known and sufficiently habituated to make strong <strong>in</strong>ference about her mat<strong>in</strong>g<br />

behavior. Follow<strong>in</strong>g the 1998 <strong>in</strong>fanticide, 3 females without dependent<br />

<strong>of</strong>fspr<strong>in</strong>g immigrated to Kasekela, presumably from Kalande (Sifa and Nasa<br />

<strong>in</strong> September and October <strong>of</strong> 2000, respectively, and Malaika <strong>in</strong> September<br />

2001). Their ages are uncerta<strong>in</strong>, though Malaika and Nasa appeared to be<br />

young adolescents that might have immigrated <strong>in</strong> any case, rather than mothers<br />

<strong>of</strong> <strong>in</strong>fanticide victims.<br />

Second, the rival coalition reduction hypothesis predicts that <strong>in</strong>tergroup<br />

attacks on both <strong>in</strong>fants and adults should be biased toward males. The <strong>in</strong>tergroup<br />

kill<strong>in</strong>g <strong>of</strong> Rejea underm<strong>in</strong>es the previously reported male bias <strong>in</strong><br />

<strong>in</strong>fanticide victims. Determ<strong>in</strong><strong>in</strong>g the sex <strong>of</strong> an <strong>in</strong>fant victim dur<strong>in</strong>g the frenzy<br />

<strong>of</strong> an <strong>in</strong>tergroup encounter is difficult. Wilson and Wrangham (2003) found<br />

that the sex <strong>of</strong> 8 <strong>of</strong> 15 <strong>in</strong>tergroup <strong>in</strong>fanticide victims is unknown. For the<br />

7 victims whose sex was determ<strong>in</strong>ed, 5 were male and 2 were female, a<br />

proportion not statistically different from the expected proportion <strong>of</strong> 50%<br />

(b<strong>in</strong>omial test, NS), which suggests that the apparent male bias is a result<br />

<strong>of</strong> small sample size, much as has been found for <strong>in</strong>tracommunity <strong>in</strong>fanti-

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