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      Selection based on the size of the black tie of the great tit may be reversed in urban habitats

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          A standard approach to model how selection shapes phenotypic traits is the analysis of capture–recapture data relating trait variation to survival. Divergent selection, however, has never been analyzed by the capture–recapture approach. Most reported examples of differences between urban and nonurban animals reflect behavioral plasticity rather than divergent selection. The aim of this paper was to use a capture–recapture approach to test the hypothesis that divergent selection can also drive local adaptation in urban habitats. We focused on the size of the black breast stripe (i.e., tie width) of the great tit ( Parus major), a sexual ornament used in mate choice. Urban great tits display smaller tie sizes than forest birds. Because tie size is mostly genetically determined, it could potentially respond to selection. We analyzed capture/recapture data of male great tits in Barcelona city ( N = 171) and in a nearby (7 km) forest ( N = 324) from 1992 to 2008 using MARK. When modelling recapture rate, we found it to be strongly influenced by tie width, so that both for urban and forest habitats, birds with smaller ties were more trap-shy and more cautious than their larger tied counterparts. When modelling survival, we found that survival prospects in forest great tits increased the larger their tie width (i.e., directional positive selection), but the reverse was found for urban birds, with individuals displaying smaller ties showing higher survival (i.e., directional negative selection). As melanin-based tie size seems to be related to personality, and both are heritable, results may be explained by cautious personalities being favored in urban environments. More importantly, our results show that divergent selection can be an important mechanism in local adaptation to urban habitats and that capture–recapture is a powerful tool to test it.

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          Most cited references 76

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          Ecological Speciation

           Patrik Nosil (2012)
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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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              The ecology of adaptive radiation


                Author and article information

                Ecol Evol
                Ecol Evol
                Ecology and Evolution
                BlackWell Publishing Ltd (Oxford, UK )
                July 2014
                07 June 2014
                : 4
                : 13
                : 2625-2632
                [1 ]Natural History Museum of Barcelona Barcelona, Spain
                [2 ]Warnell School of Forestry and Natural Resources, University of Georgia Athens, Georgia
                [3 ]Department of Ecology and Evolution, Evolutionary Biology Centre, University of Uppsala Uppsala, Sweden
                Author notes
                Juan Carlos Senar, Evolutionary and Behavioural Ecology Associate Research Unit, CSIC, Natural History Museum of Barcelona, P. Picasso s/n, E-08003 Barcelona, Spain. Tel: +34 932562216; Fax: +34 933104999; E-mail: jcsenar@

                Funding Information The present study was funded by CGL2012-38262 research project to JCS & JQ and by the FPI grants FP2000-6439 to JQ and BES2007-16320 to FM, from the Spanish Research Council (Ministry of Science and Innovation and the Ministry of Economics and Competiveness).

                © 2014 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.

                This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

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