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      Macroevolutionary patterning of woodpecker drums reveals how sexual selection elaborates signals under constraint

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          Abstract

          Sexual selection drives elaboration in animal displays used for competition and courtship, but this process is opposed by morphological constraints on signal design. How do interactions between selection and constraint shape display evolution? One possibility is that sexual selection continues exaggeration under constraint by operating differentially on each signal component in complex, modular displays. This is seldom studied on a phylogenetic scale, but we address the issue herein by studying macroevolutionary patterning of woodpecker drum displays. These territorial displays are produced when an individual rapidly hits its bill on a hard surface, and drums vary across species in the number of beats included (length) and the rate of drumbeat production (speed). We report that species body size limits drum speed, but not drum length. As a result of this biomechanical constraint, there is less standing variation in speed than length. We also uncover a positive relationship between sexual size dimorphism and the unconstrained trait (length), but with no effect on speed. This suggests that when morphology limits the exaggeration of one component, sexual selection instead exaggerates the unconstrained trait. Modular displays therefore provide the basis for selection to find novel routes to phenotypic elaboration after previous ones are closed.

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          Controlling the False Discovery Rate: A Practical and Powerful Approach to Multiple Testing

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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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              Size-correction and principal components for interspecific comparative studies.

              Phylogenetic methods for the analysis of species data are widely used in evolutionary studies. However, preliminary data transformations and data reduction procedures (such as a size-correction and principal components analysis, PCA) are often performed without first correcting for nonindependence among the observations for species. In the present short comment and attached R and MATLAB code, I provide an overview of statistically correct procedures for phylogenetic size-correction and PCA. I also show that ignoring phylogeny in preliminary transformations can result in significantly elevated variance and type I error in our statistical estimators, even if subsequent analysis of the transformed data is performed using phylogenetic methods. This means that ignoring phylogeny during preliminary data transformations can possibly lead to spurious results in phylogenetic statistical analyses of species data.
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                Author and article information

                Journal
                Proc Biol Sci
                Proc. Biol. Sci
                RSPB
                royprsb
                Proceedings of the Royal Society B: Biological Sciences
                The Royal Society
                0962-8452
                1471-2954
                28 February 2018
                21 February 2018
                21 February 2018
                : 285
                : 1873
                Affiliations
                Department of Biology, Wake Forest University , Winston-Salem, NC 27101, USA
                Author notes

                Electronic supplementary material is available online at https://dx.doi.org/10.6084/m9.figshare.c.3999561.

                Article
                rspb20172628
                10.1098/rspb.2017.2628
                5832706
                29467264
                cc4a4e97-29ab-4879-859e-fa3a643306ac
                © 2018 The Authors.

                Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original author and source are credited.

                Product
                Funding
                Funded by: National Science Foundation, http://dx.doi.org/10.13039/100000001;
                Award ID: IOS-1655730
                Categories
                1001
                14
                70
                Evolution
                Research Article
                Custom metadata
                February 28, 2018

                Life sciences
                sexual selection,animal behaviour,signal design,constraint
                Life sciences
                sexual selection, animal behaviour, signal design, constraint

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