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      The evolution of male mate choice and female ornamentation: a review of mathematical models

      research-article
      1 , 2 , Handling editor: Ingo Schlupp
      Current Zoology
      Oxford University Press
      female ornaments, male mate preferences, mathematical models

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          Abstract

          The evolution of male preferences and of female ornaments in species with traditional sex roles (i.e., polygyny) have been highlighted as areas in need of more active research by an accumulation of recent findings. The theoretical literature on these topics is relatively small and has centered on the evolution of male choice. Mathematical models have emphasized that, under polygyny, the evolution of male preferences faces much greater competition costs than does the evolution of female preferences. We discuss ways in which costly male choice can nonetheless evolve, via (1) direct selection that favors preferences, primarily through mating with highly fecund females, (2) mechanisms that rely on indirect selection, which weakly counters competitive costs of male preferences, and (3) genetic constraints, primarily in the form of pleiotropy of male and female preferences and traits. We also review a variety of mathematical models that have elucidated how costs to male preferences can be avoided. Finally, we turn our attention to the relatively scant theoretical literature on the effects of male mate choice on the evolution of female traits. We emphasize the finding that the presence of male preferences cannot be assumed to lead to the evolution of female ornaments during polygyny, and point out situations where models have elucidated ways in which female ornaments can nevertheless evolve.

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          The evolution of male mate choice in insects: a synthesis of ideas and evidence.

          Mate choice by males has been recognized at least since Darwin's time, but its phylogenetic distribution and effect on the evolution of female phenotypes remain poorly known. Moreover, the relative importance of factors thought to underlie the evolution of male mate choice (especially parental investment and mate quality variance) is still unresolved. Here I synthesize the empirical evidence and theory pertaining to the evolution of male mate choice and sex role reversal in insects, and examine the potential for male mating preferences to generate sexual selection on female phenotypes. Although male mate choice has received relatively little empirical study, the available evidence suggests that it is widespread among insects (and other animals). In addition to 'precopulatory' male mate choice, some insects exhibit 'cryptic' male mate choice, varying the amount of resources allocated to mating on the basis of female mate quality. As predicted by theory, the most commonly observed male mating preferences are those that tend to maximize a male's expected fertilization success from each mating. Such preferences tend to favour female phenotypes associated with high fecundity or reduced sperm competition intensity. Among insect species there is wide variation in mechanisms used by males to assess female mate quality, some of which (e.g. probing, antennating or repeatedly mounting the female) may be difficult to distinguish from copulatory courtship. According to theory, selection for male choosiness is an increasing function of mate quality variance and those reproductive costs that reduce, with each mating, the number of subsequent matings that a male can perform ('mating investment') Conversely, choosiness is constrained by the costs of mate search and assessment, in combination with the accuracy of assessment of potential mates and of the distribution of mate qualities. Stronger selection for male choosiness may also be expected in systems where female fitness increases with each copulation than in systems where female fitness peaks at a small number of matings. This theoretical framework is consistent with most of the empirical evidence. Furthermore, a variety of observed male mating preferences have the potential to exert sexual selection on female phenotypes. However, because male insects typically choose females based on phenotypic indicators of fecundity such as body size, and these are usually amenable to direct visual or tactile assessment, male mate choice often tends to reinforce stronger vectors of fecundity or viability selection, and seldom results in the evolution of female display traits. Research on orthopterans has shown that complete sex role reversal (i.e. males choosy, females competitive) can occur when male parental investment limits female fecundity and reduces the potential rate of reproduction of males sufficiently to produce a female-biased operational sex ratio. By contrast, many systems exhibiting partial sex role reversal (i.e. males choosy and competitive) are not associated with elevated levels of male parental investment, reduced male reproductive rates, or reduced male bias in the operational sex ratio. Instead, large female mate quality variance resulting from factors such as strong last-male sperm precedence or large variance in female fecundity may select for both male choosiness and competitiveness in such systems. Thus, partial and complete sex role reversal do not merely represent different points along a continuum of increasing male parental investment, but may evolve via different evolutionary pathways.
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            Why are female birds ornamented?

            Sexual selection is now widely accepted as the main evolutionary explanation of extravagant male ornaments. By contrast, ornaments occurring in females have received little attention and often have been considered as nonadaptive, correlated effects of selection on males. However, recent comparative evidence suggests that female ornaments have evolved quite independently of male showiness. Also, new theoretical models predict that both male mate choice and female contest competition will occur under certain circumstances. This is supported by recent experimental studies. Thus, selection acting on females might be a widespread cause of female ornaments.
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              Armaments and ornaments: an evolutionary explanation of traits of dual utility

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                Author and article information

                Journal
                Curr Zool
                Curr Zool
                czoolo
                Current Zoology
                Oxford University Press
                1674-5507
                2396-9814
                June 2018
                18 April 2018
                18 April 2018
                : 64
                : 3
                : 323-333
                Affiliations
                [1 ]Department of Biology, Indiana University, Bloomington, IN 47405, USA
                [2 ]Department of Biology, University of North Carolina at Chapel Hill, Chapel Hill, NC 27516, USA
                Author notes
                Address correspondence to Courtney L. Fitzpatrick. E -mail: cfitzpa@ 123456indiana.edu .
                Article
                zoy029
                10.1093/cz/zoy029
                6007321
                30402075
                057ab73a-e2c6-4006-b42b-8ab2222671da
                © The Author(s) (2018). Published by Oxford University Press.

                This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License ( http://creativecommons.org/licenses/by-nc/4.0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited. For commercial re-use, please contact journals.permissions@oup.com

                History
                : 28 October 2017
                : 29 March 2018
                Page count
                Pages: 11
                Funding
                Funded by: National Institutes of Health 10.13039/100000002
                Award ID: 5T32HD049336-12
                Categories
                Special Column: Male mate choice, female competition, and female ornaments
                Guest Editor: Ingo Schlupp, Department of Biology, University of Oklahoma, USA
                Articles

                female ornaments,male mate preferences,mathematical models

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