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      A dominant allele controls development into female mimic male and diminutive female ruffs

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          Abstract

          Maintaining polymorphisms for genes with effects of ecological significance may involve conflicting selection in males and females. We present data from a captive population of ruffs ( Philomachus pugnax) showing that a dominant allele controls development into both small, ‘female mimic’ males (‘faeders’), and a previously undescribed class of small ‘female faeders’. Most male ruffs have elaborate breeding plumage and display behaviour, but 0.5–1.5% are faeders, which lack both. Females from a captive population previously lacking faeders were bred with two founder faeder males and their faeder sons. The faeders’ offspring had a quadrimodal size distribution comprising normal-sized males and females, faeders and atypically small females. By contrast, ornamented males fathered only normal-sized offspring. We conclude that both founding faeders were heterozygous for a faeder allele absent from the original population. This allele is dominant to previously described genes that determine development into independent versus satellite ornamented males. Unlike those genes, the faeder allele is clearly expressed in females. Small body size is a component of the male faeder mating strategy, but provides no obvious benefit to females. Bisexual expression of the gene provides the opportunity to quantify the strength of sexually antagonistic selection on a Mendelian trait.

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          Most cited references4

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          Intralocus sexual conflict.

          Intralocus sexual conflict occurs when selection on a shared trait in one sex displaces the other sex from its phenotypic optimum. It arises because many shared traits have a common genetic basis but undergo contrasting selection in the sexes. A recent surge of interest in this evolutionary tug of war has yielded evidence of such conflicts in laboratory and natural populations. Here we highlight outstanding questions about the causes and consequences of intralocus sexual conflict at the genomic level, and its long-term implications for sexual coevolution. Whereas recent thinking has focussed on the role of intralocus sexual conflict as a brake on sexual coevolution, we urge a broader appraisal that also takes account of its potential to drive adaptive evolution and speciation.
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            The status of the conditional evolutionarily stable strategy.

            The conditional evolutionarily stable strategy (ESS) has proven to be a versatile tool for understanding the production of alternative phenotypes in response to environmental cues. Hence, we would expect the theoretical basis of the conditional strategy to be robust. However, Shuster and Wade have recently criticized the conditional ESS based on Gross's 1996 proposal that most alternative reproductive tactics are conditional and have evolved by 'status-dependent selection.' We critically assess Gross's status-dependent selection model and Shuster and Wade's critique. We find shortcomings and misconceptions in both. We return to the findings of the strategic models behind the conditional ESS and demonstrate how environmental threshold models use a reaction norm approach and quantitative genetic theory to understand the evolution of conditional strategies.
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              Permanent female mimics in a lekking shorebird.

              Female mimics are known from many species, but permanent, non-conditional, alternative mating strategies are only known from an isopod, a fish, a lizard and a bird. The single bird example refers to lek-breeding ruffs Philomachus pugnax, a shorebird for which two strategies (independent and satellite) have been known for over 50 years. Ruffs also provided the single case of an animal with two, rather than three, permanent alternative mating strategies. Here, we describe a rare female-like morph of ruffs: the 'missing' third alternative mating strategy, which we have called 'faeder'. Faeders are slightly larger than females and in late April have testes 2.5 time the size of testes of normal males. On leks in aviaries and in the wild they appear to combine feminine and masculine behaviours. Faeders may represent the ancestral, care-giving, male strategy, but their relatively large testes suggest that currently they behave as sneakers.
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                Author and article information

                Journal
                Biol Lett
                Biol. Lett
                RSBL
                roybiolett
                Biology Letters
                The Royal Society
                1744-9561
                1744-957X
                23 December 2013
                23 December 2013
                : 9
                : 6
                : 20130653
                Affiliations
                [1 ]Evolutionary Behavioural Ecology Research Group, Simon Fraser University , Burnaby, British Columbia, CanadaV5A 1S6
                [2 ]Department of Animal and Plant Sciences, University of Sheffield , Sheffield S10 2TN, UK
                [3 ]Chair in Global Flyway Ecology, Animal Ecology Group, Centre for Ecological and Evolutionary Studies, University of Groningen , PO Box 11103, 9700 CC Groningen, The Netherlands
                [4 ]Department of Marine Ecology, Royal Netherlands Institute for Sea Research (NIOZ) , PO Box 59, 1790 AB Den Burg, Texel, The Netherlands
                [5 ]Department of Biology and Center for Biodiversity, East Carolina University , Greenville, NC 27858-4353, USA
                Author notes
                Article
                rsbl20130653
                10.1098/rsbl.2013.0653
                3871350
                24196515
                a1fdb7ca-942c-406d-b845-e6c14529f7c0

                © 2013 The Authors. Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0/, which permits unrestricted use, provided the original author and source are credited.

                History
                : 19 July 2013
                : 14 October 2013
                Categories
                14
                1001
                70
                Animal Behaviour
                Custom metadata
                December 23, 2013

                Life sciences
                polymorphism,mendelian genetics,alternative male strategies,philomachus pugnax,female mimic

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