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      Adaptive coloration in pied flycatchers ( Ficedula hypoleuca)—The devil is in the detail

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          Abstract

          Understanding the origin and persistence of phenotypic variation within and among populations is a major goal in evolutionary biology. However, the eagerness to find unadulterated explanatory models in combination with difficulties in publishing replicated studies may lead to severe underestimations of the complexity of selection patterns acting in nature. One striking example is variation in plumage coloration in birds, where the default adaptive explanation often is that brightly colored individuals signal superior quality across environmental conditions and therefore always should be favored by directional mate choice. Here, we review studies on the proximate determination and adaptive function of coloration traits in male pied flycatchers ( Ficedula hypoleuca). From numerous studies, we can conclude that the dark male color phenotype is adapted to a typical northern climate and functions as a dominance signal in male–male competition over nesting sites, and that the browner phenotypes are favored by relaxed intraspecific competition with more dominant male collared flycatchers ( Ficedula albicollis) in areas where the two species co‐occur. However, the role of avoidance of hybridization in driving character displacement in plumage between these two species may not be as important as initially thought. The direction of female choice on male coloration in pied flycatchers is not simply as opposite in direction in sympatry and allopatry as traditionally expected, but varies also in relation to additional contexts such as climate variation. While some of the heterogeneity in the observed relationships between coloration and fitness probably indicate type 1 errors, we strongly argue that environmental heterogeneity and context‐dependent selection play important roles in explaining plumage color variation in this species, which probably also is the case in many other species studied in less detail.

          Abstract

          We review numerous studies on both proximate and ultimate factors behind extensive plumage color variation in male pied flycatchers and particularly zoom in on the most unknown questions and suggest future research avenues. There is strong evidence that interspecific interactions with collared flycatchers are of crucial importance for explaining the origin and maintenance of plumage color variation in male pied flycatchers, but the role of avoidance of hybridization in driving character displacement in plumage between the two flycatcher species may not be as important as initially thought. Our review suggests that plumage traits should not be expected to signal high quality as such but rather different male competitive strategies, and a main take‐home message is that selection patterns acting on signaling traits are more diverse and fluctuating than generally expected.

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          Biological signals as handicaps.

          An ESS model of Zahavi's handicap principle is constructed. This allows a formal exposition of how the handicap principle works, and shows that its essential elements are strategic. The handicap model is about signalling, and it is proved under fairly general conditions that if the handicap principle's conditions are met, then an evolutionarily stable signalling equilibrium exists in a biological signalling system, and that any signalling equilibrium satisfies the conditions of the handicap principle. Zahavi's major claims for the handicap principle are thus vindicated. The place of cheating is discussed in view of the honesty that follows from the handicap principle. Parallel signalling models in economics are discussed. Interpretations of the handicap principle are compared. The models are not fully explicit about how females use information about male quality, and, less seriously, have no genetics. A companion paper remedies both defects in a model of the handicap principle at work in sexual selection.
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            The Lek Paradox and the Capture of Genetic Variance by Condition Dependent Traits

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              Pleiotropy in the melanocortin system, coloration and behavioural syndromes.

              In vertebrates, melanin-based coloration is often associated with variation in physiological and behavioural traits. We propose that this association stems from pleiotropic effects of the genes regulating the synthesis of brown to black eumelanin. The most important regulators are the melanocortin 1 receptor and its ligands, the melanocortin agonists and the agouti-signalling protein antagonist. On the basis of the physiological and behavioural functions of the melanocortins, we predict five categories of traits correlated with melanin-based coloration. A review of the literature indeed reveals that, as predicted, darker wild vertebrates are more aggressive, sexually active and resistant to stress than lighter individuals. Pleiotropic effects of the melanocortins might thus account for the widespread covariance between melanin-based coloration and other phenotypic traits in vertebrates.
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                Author and article information

                Contributors
                paivi.sirkia@gmail.com
                Journal
                Ecol Evol
                Ecol Evol
                10.1002/(ISSN)2045-7758
                ECE3
                Ecology and Evolution
                John Wiley and Sons Inc. (Hoboken )
                2045-7758
                24 January 2021
                February 2021
                : 11
                : 4 ( doiID: 10.1002/ece3.v11.4 )
                : 1501-1525
                Affiliations
                [ 1 ] Finnish Museum of Natural History Zoology Unit University of Helsinki Helsinki Finland
                [ 2 ] Department of Ecology and Genetics Animal Ecology Uppsala University Uppsala Sweden
                Author notes
                [*] [* ] Correspondence

                Päivi M. Sirkiä, Finnish Environment Institute, Latokartanonkaari 11, 00790 Helsinki, Finland.

                Email: paivi.sirkia@ 123456gmail.com

                Author information
                https://orcid.org/0000-0003-0346-6298
                https://orcid.org/0000-0002-1178-4053
                Article
                ECE37048
                10.1002/ece3.7048
                7882974
                33613985
                ee127ec0-08cc-480c-b395-4e5162e48d47
                © 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.

                This is an open access article under the terms of the http://creativecommons.org/licenses/by/4.0/ License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

                History
                : 09 June 2020
                : 22 October 2020
                : 28 October 2020
                Page count
                Figures: 4, Tables: 3, Pages: 25, Words: 20562
                Funding
                Funded by: Academy of Finland , open-funder-registry 10.13039/501100002341;
                Award ID: 267430
                Funded by: Emil Aaltosen Säätiö , open-funder-registry 10.13039/501100004756;
                Categories
                Review Article
                Review Article
                Custom metadata
                2.0
                February 2021
                Converter:WILEY_ML3GV2_TO_JATSPMC version:5.9.7 mode:remove_FC converted:15.02.2021

                Evolutionary Biology
                ficedula hypoleuca,melanin coloration,ornaments,pied flycatcher,plumage coloration,sexual selection

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