Abstract

Male coalition-like formation, recently found in stump-tailed macaques (Macaca arctoides), occurs when several top-ranking males collaboratively guard females to prevent mating with other rival males and actively share secured mating opportunities with their allies. We lack a comprehensive understanding of the proximate mechanisms underlying such male coalitions, e.g., the effect of genetic relatedness. Such cooperative partner choice among males is particularly interesting in animals, such as M. arctoides, that live in matrilineal (female philopatry/male dispersal) society. Theoretically, in such a social system, females, and not males, are often related to each other by kinship. Thus, the occurrence of cooperative behavior between related females are generally common, but it would be rare between unrelated/non-kin males in such matrilineal society. Herein, for the first time, we report detailed copulatory behaviors, including mating and reproductive success, in relation to male coalition-like formation in free-ranging M. arctoides following direct behavioral observation and genetic analysis. We found that coalition-forming male–male alliances often occur among both individuals that are highly related and those that are less related. We identified two groups with different mating strategies, i.e., single-male monopoly and coalition-male monopoly groups. In both groups, nearly 80% of copulations were monopolized by a single male or by coalition males. However, the single-male monopoly strategy allows opportunistic/sneaky copulations by other males with a relatively high probability. Thus, the degree of reproductive success did not reflect mating success. In contrast, the males employing a coalition strategy successfully shared their mating and reproductive success, particularly in the largest group. Compared with single-male monopolized groups, the coalition-male monopoly groups copulated with a considerably more number of females, suggesting that coalition males can effectively guard against opportunistic/sneaky copulation by rival males. We also found that coalition-forming male–male alliances often occur regardless of the degree of kinship/relatedness, indicating the complexity and flexibility inherent in the male social bond of M. arctoides.

Highlights

  • A fundamental problem in behavioral ecology is determining mechanisms that regulate differential reproduction among individuals, and intrasexual competition is considered one of the important factors closely associated with such mechanisms (Darwin, 1871)

  • Male reproductive success has been studied in many animal taxa to understand differential reproduction in males possibly caused by male–male competition (Clutton-Brock, 1988)

  • Among primates that exhibit a stunning variation in both social structure and organization, male reproductive success has been historically difficult to measure, and male mating success is assumed to be a sufficiently reliable indicator to reflect male reproductive success, wherein success is often linked to dominance status (Robinson, 1982; Fedigan, 1983; Cowlishaw and Dunbar, 1991)

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Summary

Introduction

A fundamental problem in behavioral ecology is determining mechanisms that regulate differential reproduction among individuals, and intrasexual competition is considered one of the important factors closely associated with such mechanisms (Darwin, 1871). Male reproductive success has been studied in many animal taxa to understand differential reproduction (or reproductive skew) in males possibly caused by male–male competition (Clutton-Brock, 1988). Among primates that exhibit a stunning variation in both social structure and organization, male reproductive success has been historically difficult to measure, and male mating success is assumed to be a sufficiently reliable indicator to reflect male reproductive success, wherein success is often linked to dominance status (Robinson, 1982; Fedigan, 1983; Cowlishaw and Dunbar, 1991). Some studies show that mating and reproductive success are both closely related to male dominance status (e.g., Cowlishaw and Dunbar, 1991; de Ruiter and van Hooff, 1993; Wickings et al, 1993), the skew in male reproductive success is often less pronounced than that predicted from observed mating success (e.g., Berard et al, 1993; Inoue et al, 1993; Constable et al, 2001; Young et al, 2013). The variation in the relationship between mating and reproductive success is increasingly drawing attention

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