Abstract
In primates, females typically drive the evolution of the social system and present a wide diversity of social structures. To understand this diversity, it is necessary to document the consistency and/or flexibility of female social structures across and within species, contexts, and environments. Macaques (Macaca sp.) are an ideal taxon for such comparative study, showing both consistency and variation in their social relations. Their social styles, constituting robust sets of social traits, can be classified in four grades, from despotic to tolerant. However, tolerant species are still understudied, especially in the wild. To foster our understanding of tolerant societies and to assess the validity of the concept of social style, we studied female crested macaques, Macaca nigra, under entirely natural conditions. We assessed their degree of social tolerance by analyzing the frequency, intensity, and distribution of agonistic and affiliative behaviors, their dominance gradient, their bared-teeth display, and their level of conciliatory tendency. We also analyzed previously undocumented behavioral patterns in grade 4 macaques: reaction upon approach and distribution of affiliative behavior across partners. We compared the observed patterns to data from other populations of grade 4 macaques and from species of other grades. Overall, female crested macaques expressed a tolerant social style, with low intensity, frequently bidirectional, and reconciled conflicts. Dominance asymmetry was moderate, associated with an affiliative bared-teeth display. Females greatly tolerated one another in close proximity. The observed patterns matched the profile of other tolerant macaques and were outside the range of patterns of more despotic species. This study is the first comprehensive analysis of females’ social behavior in a tolerant macaque species under natural conditions and as such, contributes to a better understanding of macaque societies. It also highlights the relevance of the social style concept in the assessment of the degree of tolerance/despotism in social systems. Am. J. Primatol. 75:361-375, 2013. © 2013 Wiley Periodicals, Inc.
Highlights
In nonhuman primates, females typically drive the evolution of the social system, highlighting the importance of focusing on females when studying primate social evolution [Clutton-Brock & Lukas, 2012; Lindenfors et al, 2004]
To extend our knowledge of social behaviors of tolerant macaque species in the wild, we studied female social behaviors in two wild, habituated but not provisioned groups of crested macaque, M. nigra [Fooden, 1969], a member of grade 4, in Sulawesi, Indonesia
Crested macaques are endemic to the island of Sulawesi, Indonesia [Sugardjito et al, 1989]
Summary
Females typically drive the evolution of the social system, highlighting the importance of focusing on females when studying primate social evolution [Clutton-Brock & Lukas, 2012; Lindenfors et al, 2004]. Previous studies on female-bonded groups (i.e., with female philopatry) have investigated ecological factors influencing female social relationships [Koenig, 2002], they have looked at the various social trade-offs faced by females in terms of cooperation and competition One such trade-off is exemplified by the degree of social tolerance between dominant and subordinate individuals, which has shaped alternative conflict management strategies [de Waal, 1986], tightly linked to sociality, and potentially, differential access to resources, whether social or ecological [van Schaik, 1989]. There is pronounced dominance asymmetry between individuals and the silent bared-teeth display serves as a formal submissive signal [de Waal & Luttrell, 1985; Preuschoft & van Schaik, 2000] In these species, dominance rank and kinship markedly constrain social interactions [Chapais, 1983]. We expected the wild population’s behavioral profile to be consistent with captive conspecifics and other grade 4 macaque populations but substantially different from macaques of other grades
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