Social Relationships Among Wild Male Crested Macaques (Macaca nigra):
[Thesis]
Tyrrell, Maura E.
Social Style, Conflict Management and Coalitions
Berman, Carol
State University of New York at Buffalo
2021
206
Ph.D.
State University of New York at Buffalo
2021
Socioecological theory predicts that within-group competition over fertile females should prevent affiliative, tolerant and cooperative relationships among males of multimale, multifemale primate groups. The past few decades of research have revealed that there is much greater variation in male-male relationships than expected, but much of this variation is not well understood. Across species, males differ in the nature and patterning of dominant, aggressive and affiliative interactions, and in the ways they manage conflicts and cooperate with one another. Levels of within-group and between-group competition may influence these behavior patterns among males, yet there is currently no consensual explanation for the variation observed, due partly to the lack of detailed studies on male-male relationships of several species. In this dissertation I aim to fill in some of these gaps by examining the social style, conflict management and coalition patterns in wild male crested macaques (Macaca nigra) in Tangkoko Nature Reserve, Indonesia, a species in which we previously had limited and inconsistent perspectives on male-male relationships with some studies depicting them as rigidly hierarchical and antagonistic and others as highly tolerant. In Chapter 2, I characterize crested macaque male-male relationships within the framework of Thierry's structural model, which states that traits related to dominance asymmetry, aggression and conflict management are inherent, structurally inter-related and species-typical (Thierry, 2000, 2007). Specifically, species are hypothesized to fall on a continuum from extremely despotic to extremely tolerant. However, I found that crested macaques show a unique set of characteristics that defy this characterization. Compared to other macaque males they display a mix of highly despotic and highly tolerant traits. In stark contrast to their extremely tolerant female counterparts, crested macaque males are not highly affiliative nor closely bonded with one another. Although they engage in low levels of post-conflict affiliation, ritualized greetings and coalitionary support (see below), they appear to primarily avoid all types of interaction with one another. Taking their mix of tolerant and despotic traits together with their low frequencies of all types of contact, I label crested macaque male social style as "avoidant," rather than distinctly despotic, tolerant or intermediate. While post-conflict affiliation is the most common conflict management mechanism observed in primates, its precise function is not well understood and appears to vary across species and among the sexes (Arnold & Aureli, 2007). In Chapter 3 I test four functions of post-conflict affiliation among male crested macaques: stress reduction, relationship repair, integrated hypothesis (an integration of both the stress reduction and relationship repair functions) and benign intent signaling. My findings suggest that post-conflict affiliation does not appear to be a robust conflict management mechanism in general for crested macaque males, but it may serve as a mechanism that allows some dyads to build/maintain the trust necessary to engage in coalitions. Unlike many species, post-conflict affiliation in crested macaque males does not appear to reduce anxiety, signal benign intent nor repair relationships, except perhaps for dyads that support one another. Thus, although crested macaque males do not appear to have strong affiliative bonds, post-conflict affiliation involving body contact may function to maintain relationships with valuable coalition partners. Crested macaque male conflict management patterns directly relate to their avoidant social style; rather than attempt to make physical contact to "reconcile" after aggression, I hypothesize that males mainly use general avoidance to prevent conflict or the escalation of aggression once it has started. In Chapter 4 I examine coalition patterns and coalition partner choice in crested macaque males within the context of within-group and between-group competition by distinguishing between resident and outside male targets. Consistent with predictions for socioecology theory, crested macaque males appear to choose their coalition partners based on certain relationship qualities. Specifically, dyads with relatively high conciliatory tendencies and greeting rates were more likely than other dyads to engage in coalitions directed against resident males. I suggest that dyads with those qualities are able to build enough trust/tolerance to cooperate in attacks against resident rivals. In particular, the results are consistent with my tentative suggestion that greetings may be a way for males to test the willingness of their partner to engage in coalitions and/or invest in their relationship. Contrary to predictions for coalitions directed against outside males, coalition partner choice does not appear to be based on feasibility or opportunity. Partner choice in coalitions directed at outside males appears to be more relaxed and involves more male participants than coalitions directed at residents, indicating more broad-based cooperation against potential immigrants. These findings suggest that crested macaque males are able to cooperate and to display flexibility in their cooperative behavior depending on the target type in spite of low levels of affiliation. This dissertation reveals more complex patterns of social interaction in crested macaque males than their original depictions. I highlight the importance of cross-species comparisons that utilize several behavioral measures in addition to social style indicators to characterize male social style. Although post-conflict affiliation is widely assumed to be a robust mechanism for repairing affiliative and tolerant relationships, my findings suggest that it plays a more limited role in crested macaque males of building/maintaining the ability to cooperate in coalitions against resident targets, consistent with their limited affiliative interactions. They also illustrate the value of assessing a variety of functional hypotheses about post-conflict affiliation in light of the social style of the interactants, rather than assuming that it functions similarly across species. Evaluating aspects of male-male relationships involving conflict management and cooperation within the context of within-group and between-group competition can also help shed light on our current understanding of socioecology and the evolution of primate male relationships.