To determine who fathers the offspring in wild mountain gorilla groups containing more than one adult male silverback, we genotyped nearly one-fourth (n ؍ 92) of the mountain gorillas (Gorilla beringei beringei) living in the Virunga Volcanoes region of Africa. Paternity analysis of 48 offspring born into four groups between 1985 and 1999 revealed that, although all infants were sired by within-group males, the socially dominant silverback did not always monopolize reproduction within his group. Instead, the second-ranking male sired an average of 15% of group offspring. This result, in combination with previous findings that secondranking males fare best by not leaving the group but by staying and waiting to assume dominance even if no reproduction is possible while waiting, is not consistent with expectations from a reproductive skew model in which the silverback concedes controllable reproduction to the second-ranking male. Instead, the data suggest a ''tug-of-war'' scenario in which neither the dominant nor the second-ranking male has full control over his relative reproductive share. The two top-ranked males were typically unrelated and this, in combination with the mixed paternity of group offspring, means that multimale gorilla groups do not approximate family groups. Instead, as long-term assemblages of related and unrelated individuals, gorilla groups are similar to chimpanzee groups and so offer interesting possibilities for kinbiased interactions among individuals.genotyping ͉ Gorilla beringei beringei ͉ noninvasive sampling ͉ paternity ͉ reproductive skew
In recent years, extended altruism towards unrelated group members has been proposed to be a unique characteristic of human societies. Support for this proposal seemingly came from experimental studies on captive chimpanzees that showed that individuals were limited in the ways they shared or cooperated with others. This dichotomy between humans and chimpanzees was proposed to indicate an important difference between the two species, and one study concluded that “chimpanzees are indifferent to the welfare of unrelated group members”. In strong contrast with these captive studies, consistent observations of potentially altruistic behaviors in different populations of wild chimpanzees have been reported in such different domains as food sharing, regular use of coalitions, cooperative hunting and border patrolling. This begs the question of what socio-ecological factors favor the evolution of altruism. Here we report 18 cases of adoption, a highly costly behavior, of orphaned youngsters by group members in Taï forest chimpanzees. Half of the adoptions were done by males and remarkably only one of these proved to be the father. Such adoptions by adults can last for years and thus imply extensive care towards the orphans. These observations reveal that, under the appropriate socio-ecologic conditions, chimpanzees do care for the welfare of other unrelated group members and that altruism is more extensive in wild populations than was suggested by captive studies.
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