Abstract

Sexual coercion, in the form of forced copulations, is relatively frequently observed in orangutans and generally attributed to their semi-solitary lifestyle. High ecological costs of association for females may be responsible for this lifestyle and may have prevented the evolution of morphological fertility indicators (e.g., sexual swellings), which would attract (male) associates. Therefore, sexual conflict may arise not only about mating per se but also about associations, because males may benefit from associations with females to monitor their reproductive state and attempt to monopolize their sexual activities. Here, we evaluate association patterns and costs for females when associating with both males and females of two different orangutan species at two study sites: Suaq, Sumatra (Pongo abelii), and Tuanan, Borneo (Pongo pygmaeus wurmbii). Female association frequency with both males and females was higher in the Sumatran population, living in more productive habitat. Accordingly, we found that the cost of association, in terms of reduced feeding to moving ratio and increased time being active, is higher in the less sociable Bornean population. Males generally initiated and maintained such costly associations with females, and prolonged associations with males led to increased female fecal cortisol metabolite (FCM) levels at Tuanan, the Bornean population. We conclude that male-maintained associations are an expression of sexual conflict in orangutans, at least at Tuanan. For females, this cost of association may be responsible for the lack of sexual signaling, while needing to confuse paternity.Significance statementSocioecological theory predicts a trade-off between the benefits of sociality and the ecological costs of increased feeding competition. Orangutans’ semi-solitary lifestyle has been attributed to the combination of high association costs and low predation risk. Previous work revealed a positive correlation between association frequencies and habitat productivity, but did not measure the costs of association. In this comparative study, we show that females likely incur costs from involuntary, male-maintained associations, especially when they last for several days and particularly in the population characterized by lower association frequencies. Association maintenance therefore qualifies as another expression of sexual conflict in orangutans, and especially prolonged, male-maintained associations may qualify as an indirect form of sexual coercion.

Highlights

  • In most mammals, female reproductive success is limited by access to food resources, while that of males is mainly limited by access to females (Darwin 1871; Emlen and Oring 1977)

  • We evaluated the time females spent in association with either other females or unflanged and flanged males during follow periods in separate analyses (LMM) and with the study site, zFAI, and the age of the dependent offspring as fixed effects

  • The time to sample extraction, the total number of days an individual was followed, the age of the dependent infant, an activity budget parameter, and the Fruit Availability Index (FAI) were included in the full model to control for possible confounding factors leading to fecal cortisol metabolite (FCM) changes

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Summary

Introduction

Female reproductive success is limited by access to food resources, while that of males is mainly limited by access to females (Darwin 1871; Emlen and Oring 1977). Males and females have different behavioral strategies to optimize their lifetime fitness, which may lead to sexual conflict (Trivers 1972; Parker 1979). Male-male competition for receptive females is high, which carries a high potential for sexual conflict (Trivers 1972; Parker 1979). The relative importance of male-male competition, female choice, and sexual conflict in orangutans remains incompletely understood (Nadler 1981; Fox 1998, 2002; Knott 2009; Utami Atmoko et al 2009; Knott et al 2010; Spillmann et al 2010, 2017). Apparent physical injuries resulting from forced copulations have not been reported, and males seem to use only as much force as is necessary to achieve intromission (Knott 2009)

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