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Why the Weird Wings? University of Montana ScholarWorks at University of Montana Graduate Student Theses, Dissertations, & Graduate School Professional Papers 2019 WHY THE WEIRD WINGS? Investigating the morphology, function, and evolution of unusual feathers in pigeons and doves (Columbidae) Robert Lawrence Niese University of Montana, Missoula Let us know how access to this document benefits ouy . Follow this and additional works at: https://scholarworks.umt.edu/etd Recommended Citation Niese, Robert Lawrence, "WHY THE WEIRD WINGS? Investigating the morphology, function, and evolution of unusual feathers in pigeons and doves (Columbidae)" (2019). Graduate Student Theses, Dissertations, & Professional Papers. 11406. https://scholarworks.umt.edu/etd/11406 This Dissertation is brought to you for free and open access by the Graduate School at ScholarWorks at University of Montana. It has been accepted for inclusion in Graduate Student Theses, Dissertations, & Professional Papers by an authorized administrator of ScholarWorks at University of Montana. For more information, please contact [email protected]. WHY THE WEIRD WINGS? Investigating the morphology, function, and evolution of unusual feathers in pigeons and doves (Columbidae) By ROBERT LAWRENCE NIESE B.Sc. Biology, University of Puget Sound, Tacoma, WA, 2011 Dissertation presented in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Organismal Biology, Ecology, and Evolution The University of Montana Missoula, MT Approved by: Dr. Bret W. Tobalske, Chairperson Division of Biological Sciences, University of Montana Dr. Christopher J. Clark Department of Biology, University of California - Riverside Dr. Erick Greene Division of Biological Sciences, University of Montana Dr. Douglas J. Emlen Division of Biological Sciences, University of Montana Dr. H. Arthur Woods Division of Biological Sciences, University of Montana Dr. Scott Whittenburg, Graduate School Dean, University of Montana i TABLE OF CONTENTS OVERVIEW .......................................................................................................... 1 FIGURE 1 ............................................................................................................................ 7 FIGURE 2 ............................................................................................................................ 9 REFERENCES ..................................................................................................................... 10 CHAPTER 1 ........................................................................................................ 13 INTRODUCTION ................................................................................................................ 15 MATERIALS AND METHODS .............................................................................................. 19 RESULTS ........................................................................................................................... 25 DISCUSSION ..................................................................................................................... 29 FIGURE 1 .......................................................................................................................... 37 FIGURE 2 .......................................................................................................................... 39 FIGURE 3 .......................................................................................................................... 41 FIGURE 4 .......................................................................................................................... 42 REFERENCES ..................................................................................................................... 44 SUPPLEMENTAL MATERIAL .............................................................................................. 47 CHAPTER 2 ........................................................................................................ 53 INTRODUCTION ................................................................................................................ 55 RESULTS AND DISCUSSION ............................................................................................... 59 MATERIALS AND METHODS .............................................................................................. 65 FIGURE 1 .......................................................................................................................... 73 FIGURE 2 .......................................................................................................................... 75 FIGURE 3 .......................................................................................................................... 77 FIGURE 4 .......................................................................................................................... 78 FIGURE 5 .......................................................................................................................... 79 FIGURE 6 .......................................................................................................................... 82 SUPPLEMENTAL TABLE 1 .................................................................................................. 85 REFERENCES ..................................................................................................................... 86 CHAPTER 3 ........................................................................................................ 92 INTRODUCTION ................................................................................................................ 94 MATERIALS AND METHODS .............................................................................................. 98 RESULTS ......................................................................................................................... 103 ii DISCUSSION ................................................................................................................... 107 FIGURE 1 ........................................................................................................................ 111 FIGURE 2 ........................................................................................................................ 113 FIGURE 3 ........................................................................................................................ 114 FIGURE 4 ........................................................................................................................ 115 FIGURE 5 ........................................................................................................................ 117 FIGURE 6 ........................................................................................................................ 118 FIGURE 7 ........................................................................................................................ 119 SUPPLEMENTAL MEDIA FILES: ........................................................................................ 120 REFERENCES ................................................................................................................... 122 APPENDIX A .................................................................................................... 127 FIGURE 1 ........................................................................................................................ 129 FIGURE 3 ........................................................................................................................ 133 iii OVERVIEW Feathers are the most phenotypically diverse (Prum and Brush, 2002; Stoddard and Prum, 2011) and structurally complex (Feo et al., 2015; Prum and Williamson, 2001) integumentary structures in vertebrates. They vary in size, shape, color, structure, and chemical composition between species, between locations on the body, and even within a single feather follicle across an individual’s lifetime (Lucas and Stettenheim, 1972). However, some feathers are morphologically conserved and show comparatively little variation across disparate taxa, individual lifespans, and evolutionary time. Remiges, the feathers that make up the majority of the surface area of a wing, are constrained and consequently morphologically stereotyped by the physical demands of flight. This morphological stereotyping by selection on flight ability has shaped the avian Bauplan since the Jurassic, and paleornithologists have used the presence of these specialized, asymmetrical feathers to define the origins of flight in the fossil record (Feduccia and Tordoff, 1979). In modern birds, this morphology is required for effective and efficient flight (Hedenström, 2003), and birds go to great lengths to compensate for, and recover from the impacts of a reduction in overall wing surface area caused by feather damage or loss (DesRochers et al., 2009; Echeverry-Galvis and Hau, 2013; Hedenström, 2003; Swaddle et al., 1996). In comparison, the feathers of the tail (the rectrices), which are also typically aerodynamically specialized and asymmetrical, are released from the constraints of locomotion by “hiding in the wake” of the flying bird (Askew, 2014; Clark, 2010; Clark and Dudley, 2009). This has allowed rectrices to evolve atypical and extreme morphologies dozens, if not hundreds of times (Clark and Dudley, 2009) that function as sexually selected visual signals (e.g. 1 widowbirds, Euplectes;
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