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Geographic Variation in Rock Wren (Salpinctes Obsoletus) Song Complexity

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Geographic Variation in Rock Wren (Salpinctes Obsoletus) Song Complexity University of Northern Colorado Scholarship & Creative Works @ Digital UNC Dissertations Student Research 8-2018 Geographic Variation in Rock Wren (Salpinctes Obsoletus) Song Complexity Nadje Amal Najar Follow this and additional works at: https://digscholarship.unco.edu/dissertations Recommended Citation Najar, Nadje Amal, "Geographic Variation in Rock Wren (Salpinctes Obsoletus) Song Complexity" (2018). Dissertations. 514. https://digscholarship.unco.edu/dissertations/514 This Text is brought to you for free and open access by the Student Research at Scholarship & Creative Works @ Digital UNC. It has been accepted for inclusion in Dissertations by an authorized administrator of Scholarship & Creative Works @ Digital UNC. For more information, please contact [email protected]. © 2018 NADJE AMAL NAJAR ALL RIGHTS RESERVED UNIVERSITY OF NORTHERN COLORADO Greeley, Colorado The Graduate School GEOGRAPHIC VARIATION IN ROCK WREN (SALPINCTES OBSOLETUS) SONG COMPLEXITY A Dissertation Submitted in Partial Fulfillment Of the Requirements for the Degree of Doctor of Philosophy Nadje Amal Najar College of Natural and Health Sciences School of Biological Sciences Biological Education August 2018 This Dissertation by: Nadje Amal Najar Entitled: Geographic variation in rock wren (Salpinctes obsoletus) song complexity has been approved as meeting the requirement for the Degree of Doctor of Philosophy in the College of Natural and Health Sciences in the School of Biological Sciences, Program of Biological Education. Accepted by the Doctoral Committee _____________________________________________________ Lauryn Benedict, Ph.D., Research Advisor _____________________________________________________ Scott Franklin, Ph.D., Committee Member _____________________________________________________ Mitchell McGlaughlin, Ph.D., Committee Member _____________________________________________________ Jessica Salo, Ph.D., Faculty Representative Date of Dissertation Defense__________________________________ Accepted by the Graduate School __________________________________________________ Linda L. Black, Ed.D, Associate Provost and Dean Graduate School and International Admission ABSTRACT Najar, Nadje Amal. Geographic variation in rock wren (Salpinctes obsoletus) song complexity. Published Doctor of Education dissertation, University of Northern Colorado, 2018. Birds sing to advertise for mates and repel rivals, but there is enormous variety in how they do this. One of the best-studied and most intriguing questions in the field is how song varies in complexity from one bird to the next, at all taxonomic levels. Several studies have found associations between migratory behavior or latitudinal gradients and song complexity, but it remains unclear how universal this pattern is or what factors may be driving it. This small body of literature suffers from several problems, perhaps the most glaring of which is the lack of systematic, population-level studies. The main goals of this dissertation were to determine what evidence there is for the hypothesis that song complexity is influenced by latitude and/or migratory behavior and whether such a pattern can be detected in a single species, the rock wren (Salpinctes obsoletus). I recorded rock wren song at 11 sites in a latitudinal transect with both migratory and sedentary populations, and used morphological measurements and genome-level SNP scans to test my classification scheme of migratory versus sedentary populations. Song repertoire size was larger in sedentary rock wrens but did not vary with latitude, while iii migratory wrens had smaller mean repertoire sizes which increased with increasing latitude. Morphological measurements differed between migratory and sedentary populations, suggesting life history differences between these two groups. Population genetic structure was only apparent using outlier loci, but the resulting structure was not concordant with migratory behavior or site membership. Taken together, these results suggest migration does not pose a barrier to gene flow between migratory and sedentary populations, and that migratory and sedentary behavior is associated with differences in song complexity and morphology, although in a way inconsistent with any previously published hypotheses. iv ACKNOWLEDGEMENTS It’s been a long time coming, but we’re finally here. While it’s my name on the front page of this document, and I’ve certainly spent blood, sweat, and tears (both literal and metaphorical) assembling it, this dissertation exists as the result of the efforts of a lot of people whose contributions I am happy to acknowledge. I couldn’t have done this project without the aid of several people who helped me with the nuts and bolts of data collection and analysis. I’d like to thank the people who assisted me in the field: Charmaine Holloway, Carissa King, and Sami Piper. Dr. Mit McGlaughlin taught me all the basics of genetics lab work, and Jenna McCullough gave me a crash course in target sequence capture. Sami Naibauer helped me get started learning Linux. Dr. Garth Spellman was absolutely essential to the genetics portion of this project, and served as a sort of ghost member of my committee. He persuaded us to go with targeted sequence capture, paid for the reagents, and was my lifeline in the analysis of the genetics data, and I can’t thank him enough for volunteering to jump on board the rock wren project. Most of the people who helped me through this project did so just by being there when I needed them. To talk to, bounce ideas off of, philosophize with, rant at, or just to be shoulders to lean on, I think I’ve had interesting, existential, sanity-rescuing conversations with most of the people in the department. In particular, I’d like to thank Tom McCabe, Stephanie Pitt, Karina Sanchez, and Cara Smith, all of whom probably know as much about my project as my committee does. My discussions with them were v just as therapeutic as they were intellectual, and I’m grateful for their patience and interest. I’d also like to thank the other members of the Benedict lab, my committee, my longtime (and long suffering) partner, Jordan Rose, and my bestest birdy friends Ilse, Baudelaire, and the dozen other birds that keep me company. Finally, I’d like to thank my research advisor, Dr. Lauryn Benedict. She was the best mentor I could have had. She patiently helped me navigate the perils of graduate school, professional societies, acquiring funding, and my dissertation project. Dr. Benedict was always there to (very tactfully) poke, prod, and encourage me to go on. I’m a better student, writer, scientist, and human being as a result of her time spent mentoring me. Thank you all so, so much, Nadje vi TABLE OF CONTENTS CHAPTER I INTRODUCTION TO ANIMAL SIGNALING, SEXUAL SELECTION, BIRD SONG, AND BIRD MIGRATION………… 1 Introduction Animal Communication Signals and Sexual Selection Bird Song as a Signal Migration Study Species – Rock Wrens II THE RELATIONSHIP BETWEEN LATITUDE, MIGRATION, AND THE EVOLUTION OF BIRD SONG COMPLEXITY …… 29 Abstract Introduction Latitude, Migration, and Song Complexity Methods Results Discussion Conclusions III GEOGRAPHIC PATTERN OF ROCK WREN SONG REPERTOIRE SIZE ……………………………………………… 53 Abstract Introduction Methods Results Discussion IV MIGRATORY MORPHOLOGY AND POPULATION GENETICS OF ROCK WRENS …………………………………. 78 Abstract Introduction Methods Results Discussion vii V SYNTHESIS AND CONCLUSIONS …………………………….. 105 Song Complexity Pattern of Repertoire Size Migratory Syndrome…? Summary REFERENCES ……………………………………………………………………. 115 APPENDIX A. INSTITUTIONAL ANIMAL CARE AND USE COMMITTEE APPROVAL …………………………………………………………… 144 B. TERRITORY MAPS …………………………………………………... 146 C. TARGETED LOCI …………………………………………………….. 154 D. DISCRIMINANT ANALYSIS OF PRINCIPAL COMPONENTS POPULATION ASSIGNMENTS ……………………………………… 160 viii LIST OF TABLES CHAPTER II 2.1 Summary of studies ………………………………………….......... 33 2.2 Summary of hypotheses ………………………………………….... 37 CHAPTER III 3.1 Study sites ……….……………………………………………........ 58 3.2 Wren captures ……………………………………………………... 63 3.3 Recording effort …………………………………………………… 64 3.4 Modeling repertoire size …………………………………………... 69 3.5 Territory sizes ……………………………………………………... 69 CHAPTER IV 4.1 Capture demographics …………………………………………….. 84 4.2 Morphological measurements ……………………………………... 92 4.3 Morphological predictions ………………………………………… 92 CHAPTER V 5.1 Support for hypotheses…………………………………………….. 108 5.2 Morphological summary …………………………………………... 111 ix LIST OF FIGURES CHAPTER I 1.1 Song sparrow spectrogram ……………………………………........ 14 1.2 Songbird brain ……………………………………………………... 16 1.3 Wren distribution map …………………………………………....... 24 1.4 Rock wren songs …………………………………………………… 25 CHAPTER III 3.1 Study sites …………………………………………………….......... 58 3.2 Accumulation curves ………………………………………………. 66 3.3 Wilcoxon signed rank test …………………………………………. 67 3.4 Song scatterplot ……………………………………………………. 68 3.5 Territory map ………………………………………………………. 70 CHAPTER IV 4.1 Migratory strategies ………………………………………………... 83 4.2 Target sequence capture …………………………………………… 88 4.3 Outlier loci …………………………………………………………. 93 4.4 Population assignment likelihoods ………………………………… 94 4.5 FASTSTRUCTURE plots …………………………………………. 95 4.6 dapc assignments …………………………………………………... 97 x 1 CHAPTER I INTRODUCTION TO ANIMAL SIGNALING, SEXUAL SELECTION, BIRD SONG, AND BIRD MIGRATION Introduction Bird song has long been used to study how various selection pressures affect signal evolution, with particular emphasis on the role of sexual selection in generating signal
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