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      Dynamics of Intersexual Dominance and Adult Sex- Ratio in Wild Vervet Monkeys

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          Abstract

          Intersexual dominance relations are important for female mammals, because of their consequences for accessing food and for the degree of sexual control females experience from males. Female mammals are usually considered to rank below males in the dominance hierarchy, because of their typical physical inferiority. Yet, in some groups or species, females are nonetheless dominant over some males (partial female dominance). Intersexual dominance, therefore, also depends on traits other than sexual dimorphism, such as social support, social exchange, group adult sex-ratio, and the widespread self-reinforcing effects of winning and losing fights, the “winner-loser effect.” The importance of sex-ratio and the winner-loser effect remains poorly understood. A theoretical model, DomWorld, predicts that in groups with a higher proportion of males, females are dominant over more males when aggression is fierce (not mild). The model is based on a small number of general processes in mammals, such as grouping, aggression, the winner-loser effect, the initially greater fighting capacity of males than females, and sex ratio. We expect its predictions to be general and suggest they be examined in a great number of species and taxa. Here, we test these predictions in four groups of wild vervet monkeys ( Chlorocebus pygerythrus) in Mawana game reserve in Africa, using 7 years of data. We confirm that a higher proportion of males in the group is associated with greater dominance of females over males; a result that remains when combining these data with those of two other sites (Amboseli and Samara). We additionally confirm that in groups with a higher fraction of males there is a relatively higher (a) proportion of fights of males with other males, and (b) proportion of fights won by females against males from the fights of females with any adults. We reject alternative hypotheses that more dominance of females over males could be attributed to females receiving more coalitions from males, or females receiving lowered male aggression in exchange for sexual access (the docile male hypothesis). We conclude that female dominance relative to males is dynamic and that future empirical studies of inter-sexual dominance will benefit by considering the adult sex-ratio of groups.

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          Most cited references42

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          Modulation of aggressive behaviour by fighting experience: mechanisms and contest outcomes.

          Experience in aggressive contests often affects behaviour during, and the outcome of, later contests. This review discusses evidence for, variations in, and consequences of such effects. Generally, prior winning experiences increase, and prior losing experiences decrease, the probability of winning in later contests, reflecting modifications of expected fighting ability. We examine differences in the methodologies used to study experience effects, and the relative importance and persistence of winning and losing experiences within and across taxa. We review the voluminous, but somewhat disconnected, literature on the neuroendocrine mechanisms that mediate experience effects. Most studies focus on only one of a number of possible mechanisms without providing a comprehensive view of how these mechanisms are integrated into overt behaviour. More carefully controlled work on the mechanisms underlying experience effects is needed before firm conclusions can be drawn. Behavioural changes during contests that relate to prior experience fall into two general categories. Losing experiences decrease willingness to engage in a contest while winning experiences increase willingness to escalate a contest. As expected from the sequential assessment model of contest behaviour, experiences become less important to outcomes of contests that escalate to physical fighting.A limited number of studies indicate that integration of multiple experiences can influence current contest behaviour. Details of multiple experience integration for any species are virtually unknown. We propose a simple additive model for this integration of multiple experiences into an individual's expected fighting ability. The model accounts for different magnitudes of experience effects and the possible decline in experience effects over time. Predicting contest outcomes based on prior experiences requires an algorithm that translates experience differences into contest outcomes. We propose two general types of model, one based solely on individual differences in integrated multiple experiences and the other based on the probability contests reach the escalated phase. The difference models include four algorithms reflecting possible decision rules that convert the perceived fighting abilities of two rivals into their probabilities of winning. The second type of algorithm focuses on how experience influences the probability that a subsequent contest will escalate and the fact that escalated contests may not be influenced by prior experience. Neither type of algorithm has been systematically investigated.Finally, we review models for the formation of dominance hierarchies that assume that prior experience influences contest outcome. Numerous models have reached varied conclusions depending on which factors examined in this review are included. We know relatively little about the importance of and variation in experience effects in nature and how they influence the dynamics of aggressive interactions in social groups and random assemblages of individuals. Researchers should be very active in this area in the next decade. The role of experience must be integrated with other influences on contest outcome, such as prior residency, to arrive at a more complete picture of variations in contest outcomes. We expect that this integrated view will be important in understanding other types of interactions between individuals, such as mating and predator-prey interactions, that also are affected significantly by prior experiences.
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            Social competition and selection in males and females

            During the latter half of the last century, evidence of reproductive competition between males and male selection by females led to the development of a stereotypical view of sex differences that characterized males as competitive and aggressive, and females as passive and choosy, which is currently being revised. Here, we compare social competition and its consequences for selection in males and females and argue that similar selection processes operate in both sexes and that contrasts between the sexes are quantitative rather than qualitative. We suggest that classifications of selection based on distinction between the form of competition or the components of fitness that are involved introduce unnecessary complexities and that the most useful approach in understanding the evolution and distribution of differences and similarities between the sexes is to compare the operation of selection in males and females in different reproductive systems.
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              Towards the integration of social dominance and spatial structure.

              My aim was to show how individual-oriented (or artificial life) models may provide an integrative background for the development of theories about dominance by including effects of spatial structure. Dominance interactions are thought to serve two different, contrasting functions: acquisition of high rank and reduction of aggression. The model I present consists of a homogeneous virtual world inhabited by artificial agents whose actions are restricted to grouping and dominance interactions in which the effects of winning and losing are self-reinforcing. The two functions are implemented as strategies to initiate dominance interactions and the intensity of aggression and dominance perception (direct or memory based) are varied experimentally. Behaviour is studied by recording the same behavioural units as in real animals. Ranks appear to differentiate more clearly at high than at low intensity of aggression and also more in the case of direct than of memory-based rank perception. Strong differentiation of rank produces a cascade of unexpected effects that differ depending on which function is implemented: for instance, a decline in aggression, spatial centrality of dominants and a correlation between rank and aggression. Insight into the origination of these self-organized patterns leads to new hypotheses for the study of the social behaviour of real animals. Copyright 2000 The Association for the Study of Animal Behaviour.
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                Author and article information

                Contributors
                Journal
                Front Psychol
                Front Psychol
                Front. Psychol.
                Frontiers in Psychology
                Frontiers Media S.A.
                1664-1078
                14 May 2020
                2020
                : 11
                : 839
                Affiliations
                [1] 1Groningen Institute for Evolutionary Life Sciences, University of Groningen , Groningen, Netherlands
                [2] 2Department of Ecology and Evolution, University of Lausanne , Lausanne, Switzerland
                [3] 3Inkawu Vervet Project, Mawana Game Reserve , Kwazulu Natal, South Africa
                [4] 4Biometris, Wageningen University & Research , Wageningen, Netherlands
                Author notes

                Edited by: Sylvia Kaiser, University of Münster, Germany

                Reviewed by: Richard McFarland, University of Wisconsin–Madison, United States; Megan Petersdorf, New York University, United States

                *Correspondence: Charlotte Korinna Hemelrijk, c.k.hemelrijk@ 123456rug.nl

                These authors share last authorship

                This article was submitted to Comparative Psychology, a section of the journal Frontiers in Psychology

                Article
                10.3389/fpsyg.2020.00839
                7240123
                32477214
                4b35afa3-1bd8-422f-9b87-efdfd5406d26
                Copyright © 2020 Hemelrijk, Wubs, Gort, Botting and van de Waal.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 13 November 2019
                : 06 April 2020
                Page count
                Figures: 4, Tables: 1, Equations: 0, References: 55, Pages: 11, Words: 0
                Categories
                Psychology
                Original Research

                Clinical Psychology & Psychiatry
                the winner-loser effect,dominance hierarchy,fierceness of aggression,female dominance over males,adult sex-ratio,vervet monkeys

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