David P. Watts

Abstract

Data on a large sample of interventions that individuals in two mountain gorilla (Gorilla gorilla beringei) groups made in agonistic interactions between others corroborate and extend earlier analyses in several ways. Related females supported each other as often as those in some female-bonded primates and maintained alliances while they resided together. Most unrelated females rarely supported each other, but some developed alliances. Females mostly supported other females with whom they had affinitive relationships against those they often engaged in dyadic aggression. They showed reciprocity in support, and often intervened against individuals who, in turn, often intervened against them. But even female coalitions that outnumbered their opponents by more than two-to-one had limited effectiveness, largely because males intervened in many female contests to control aggression. By rendering coalitions ineffective, males contribute to a combination of factors (e.g. low potential to gain from cooperation in contest feeding competition) that limit the benefits of female philopatry. Male curtailment of female aggression may influence female mate choice. Co-resident mature males in the study groups competed to control female aggression. High-ranking males curtailed aggression to females by subordinates, although two males formed an alliance against two others in their group. Immature animals mostly received defensive support against larger individuals and did not receive support from adults that could lead to a nepotistic dominance system.

David P. Watts

Abstract

Male dominance hierarchies occur in many group-living primates and some non-primate mammals. Variation in aspects of agonistic relationships such as how many dyads show bidirectionality in aggression leads to variation in dominance hierarchies along a continuum from egalitarian (relatively small agonistic power differences between adjacently-ranked individuals, shallow hierarchies) to despotic (relatively large differences, steep hierarchies). Ranks usually depend mostly or entirely on individual characteristics that influence fighting ability (e.g., body size) and show inverse-U shaped relationships to age. However, coalitionary support sometimes also influences ranks. Chimpanzees (Pan troglodytes) form multi-male, multi-female communities within which males compete for status. Males typically form dominance hierarchies, and data from multiple study show that rank is positively related to paternity success. Males also often form coalitions and some dyads form long-term alliances. Effective coalitionary support can help individuals improve and maintain their ranks, and some evidence supports the hypothesis that coalitionary aggression generally, and the positions that males hold in coalitions networks specifically, influences paternity success. Hierarchy steepness varies among communities and within communities over time; variation in the number of prime-aged males per community is a likely source of this variation. Long-term data from an extremely large chimpanzee community with unusually many males, at Ngogo, Kibale National Park, Uganda, are largely consistent with previous analyses of male chimpanzee dominance hierarchies, but show several notable contrasts. Males at Ngogo formed significantly linear hierarchies and hierarchy steepness was greater than expected if the outcomes of agonistic interactions were random. However, variation in steepness did not show the significant inverse relationship to the number of “prime-aged” males documented for other chimpanzee communities and average steepness was high given the large number of males. Ranks showed an inverse-U shaped relationship to age, although individual rank trajectories varied considerably, but males attained their highest lifetime ranks at later ages and maintained relatively high ranks to later ages than those at other chimpanzee research sites. Two measures of coalition networks, strength and Bonacich power, showed significant positive relationships with male ranks. Strength is the rate at which males joined coalitions. Bonacich power is a measure of network centrality that assesses a male’s relational power, or influence (Bonacich, 1987): a male with high Bonacich power formed coalitions with relatively many other males who were also central in the coalition network, i.e., he was strongly connected to powerful others. On average, males also attained maximum values for these and other network measures relatively late and maintained relatively high values to relatively late ages. High coalition network strength, Bonacich power, and eigenvector centrality early in adulthood were associated with high peak ranks at later ages. However, the direction of causality between participation in coalition networks and ranks is not yet clear, and the effects of body size on dominance ranks and individual rank trajectories remains to be explored. Ngogo is a favourable habitat for chimpanzees and survivorship there is unusually high; this presumably facilitates the ability of males to maintain high competitive ability longer than at other sites and shifts rank trajectories toward older ages and leads to relatively steep hierarchies despite the fact that many male dyads have similar competitive ability. Future work will assess the impact of coalitions on dominance relationships in more detail and the relationship of coalitionary aggression to paternity success.

John Groeger, Julie Harris, Simon Watt, David Rose, Paul Hibbard, Tricia Riddell, Andrew Glennerster, Brian Rogers, Keith Langley and Simon Rushton