Since W. D. Hamilton's seminal work on the evolution of sociality, a large body of research has accumulated on how kin selection might explain the evolution of cooperation in many group-living species. Our study examined the evolutionary basis of philopatry and cooperation; specifically, whether individuals benefit from the presence of close kin. We applied an individual fitness approach to a 16-year study of Columbian ground squirrels (Urocitellus columbianus) to investigate potential causal paths by which the presence of kin might act on individual fitness. Our results indicate that individual fitness benefits resulted from associations of philopatric female kin, and support the hypothesis that increased tolerance of proximity of kin is a proximate mechanism for these benefits. The major life-history influence of kin on individual fitness was through improved reproductive success, and this benefit may have been owing to philopatric settlement of kin that were recognized through familiarization in the natal burrow. Thus, we demonstrated an evolutionary basis necessary for ongoing kin-selected cooperation in Columbian ground squirrels, though the mechanism of familiarity may determine which kin individuals benefit from cooperative behaviours.

Empirical and theoretical studies have supported kin selection by demonstrating nepotism or modelling its conditions and consequences. As an alternative, we previously found that female Columbian ground squirrels had greater direct fitness when more close kin were present. Extending those results, we used population matrix methods to calculate minimum estimates of individual fitness, estimated direct and indirect components of fitness, estimated inclusive fitness by adding the direct fitness (stripped of estimated influences of the social environment) and indirect fitness components together, and finally looked for inclusive fitness benefits of associations with close kin who seem to be 'genial neighbours'. We examined the estimated fitness of a sample of 35 females for which complete lifetimes were known for themselves, their mothers and their littermate sisters. Six of these females had no cosurviving adult close kin, and their direct fitness was significantly lower than 29 females with such kin (λ = 0.66 vs. λ = 1.23). The net fitness benefit of the presence of close kin was thus 0.57. The estimated indirect component of fitness through benefits to the direct fitness of close kin was 0.43. Thus, estimated inclusive fitness for females with cosurviving close kin (λ = 1.09) was significantly greater than that for females without surviving close kin (viz., λ = 0.66). The presence of closely related and philopatric female kin appeared to result in considerable fitness benefits for female ground squirrels, perhaps through the behavioural mechanisms of lowered aggression and other forms of behavioural cooperation.

To understand the evolution of personality structure requires examining personality dimensions in multiple species using a common set of traits. Little research has been conducted on personality in wild populations of nonhuman primates. Using behavioral observations and questionnaire ratings, we examined factors influencing personality in 16 wild bonobos (Pan paniscus) at Wamba, Luo Scientific Reserve, Democratic Republic of the Congo. We extracted five factors from 31 of the items from the Hominoid Personality Questionnaire (HPQ) and three factors from observed behaviors. The HPQ factors were labeled Unemotionality , Friendliness , Aggressiveness , Irritability , and Activity . The behavioral factors were labeled Grooming , Playfulness , and Introversion . We established the convergent and divergent validity of these factors by obtaining correlations between the HPQ and behavioral factors. We tested for sex differences and found that males were significantly higher on Introversion and significantly lower in Irritability . We then tested for age differences and found that Friendliness was lower and Aggressiveness was higher in older individuals. Finally, we found that, among males, hierarchical rank was associated with higher Aggressiveness . These findings contrast with findings in chimpanzees in ways consistent with known species differences. For one, consistent with the more egalitarian structure of bonobo society, we did not identify a clear Dominance factor. Also, the results related to sex differences were consistent with previous findings that reveal closer bonds between female bonobos than female chimpanzees. These findings highlight the importance of studying personality in closely related species and the need to consider species' socioecology when studying personality. Am. J. Primatol. 78:1178-1189, 2016. © 2016 Wiley Periodicals, Inc.