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Conirostrum Cinereum) Andrea N University of New Mexico UNM Digital Repository Biology ETDs Electronic Theses and Dissertations Spring 5-15-2017 Natural history and evolution of an elevational generalist, the Cinereous Conebill (Conirostrum cinereum) Andrea N. Chavez University of New Mexico Chandresekhar Natarajan University of Nebraska - Lincoln Jay F. Storz University of Nebraska - Lincoln Angela Fago Aarhus University Christopher C. Witt University of New Mexico Follow this and additional works at: https://digitalrepository.unm.edu/biol_etds Part of the Biology Commons Recommended Citation Chavez, Andrea N.; Chandresekhar Natarajan; Jay F. Storz; Angela Fago; and Christopher C. Witt. "Natural history and evolution of an elevational generalist, the Cinereous Conebill (Conirostrum cinereum)." (2017). https://digitalrepository.unm.edu/biol_etds/199 This Thesis is brought to you for free and open access by the Electronic Theses and Dissertations at UNM Digital Repository. It has been accepted for inclusion in Biology ETDs by an authorized administrator of UNM Digital Repository. For more information, please contact [email protected]. ! ! ! ! ! ! ! Andrea N. Chavez Candidate Biology Department This thesis is approved, and it is acceptable in quality and form for publication: Approved by the Thesis Committee: Christopher C. Witt, Chairperson Joseph Cook Michael Andersen ! i! NATURAL HISTORY AND EVOLUTION OF AN ELEVATIONAL GENERALIST, THE CINEREOUS CONEBILL (CONIROSTRUM CINEREUM) by ANDREA N. CHAVEZ B.S., BIOLOGY, UNIVERSITY OF NEW MEXICO, 2008 THESIS Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Science Biology The University of New Mexico Albuquerque, New Mexico May 2017 ! ii! ACKNOWLEDGMENTS I would like to first thank my major advisor, Dr. Christopher C. Witt, for the invaluable guidance and support he has provided throughout my tenure as a graduate student at the University of New Mexico. I would also like to thank my committee members, Drs. Joseph Cook and Michael Andersen, for the scientific insight and research assistance they have provided in support of my research. Pursuing a graduate degree while working as a professional wildlife biologist for a federal agency has been difficult. I greatly appreciate my committee for their patience, understanding, and support for my career goals and aspirations throughout this time. For intellectual discourse, I would like to thank Elizabeth Beckman, Ariel Gaffney, Spencer C. Galen, Andrew Johnson, Chauncey Gadek, Matthew J. Baumann, C. Jonathan Schmitt and Ashley Smiley. I would also like to thank Fernando Angulo, Emil Bautista, Daniel F. Lane, Sheila Figeroa, L. Monica Flores, Willy Ñáñez, Jano Nuñez, Alessandra Quiñonez Z., Donna C. Schmitt, Dora Susanibar, Thomas Valqui, Walter Vargas, and many associates of Centro de Ornitología y Biodiversidad (CORBIDI) for assisting with fieldwork to obtain key specimens for this research. The American Museum of Natural History Department of Ornithology graciously loaned voucher specimens that greatly improved this work. Furthermore, this research would not have been possible without funding from NSF DEB- 1146491, NSF DEB-0543556, the UNM Center for Evolutionary and Theoretical Immunology (CETI), the Graduate and Professional Student Association, the University of New Mexico Latina/o Graduate and Professional Student Fellowship, the Donald Caughran Memorial Scholarship, the Molly Hayes Scholarship and the late Robert W. Dickerman. I thank my former supervisor at the Bureau of Land Management Lori Kimball for ! iii! supporting my pursuit of a graduate degree while working as a permanent employee. I thank Marikay Ramsey for the academic advice and professional mentorship she has provided throughout my career as a wildlife biologist. I would like to especially thank Vincent Ortega for his unending support for me throughout all my educational and professional endeavors. ! iv! NATURAL HISTORY AND EVOLUTION OF AN ELEVATIONAL GENERALIST, THE CINEREOUS CONEBILL (CONIROSTRUM CINEREUM) by Andrea N. Chavez B.S., University of New Mexico, 2008 M.S., University of New Mexico, 2017 ABSTRACT Elevational generalism is relatively rare in the tropical Andes Mountains, likely owing to the inherent requirements of enduring a high degree of climatic zonation and coping with hypoxic stress. The Cinereous Conebill (Conirostrum cinereum) appears to be an exception, and inhabits a continuous elevational distribution that spans over 4,500 m. Two subspecies, cinereum and fraseri, are restricted to high elevations and may be isolated, whereas the third and most widespread, littorale, occurs continuously along the western slope of the Andes from 0 to over 4,500 m. First, we aim to characterize the morphology, genetics, and climatic niche of the three subspecies using a comparative biogeographic approach to explore patterns and timing of differentiation and to consider possible mechanisms of diversification. Second, we study whether hemoglobin adaptation plays a role in this elevational generalist’s ability to thrive in high-elevation environments, and whether localized adaptation is possible despite altitudinal migration and gene flow. We used a comparative phylogeographic framework to examine whether lineage divergence within C. cinereum is associated with climatic, geographic, and/or physiological barriers leading to incipient speciation. ! ! v! TABLE OF CONTENTS INTRODUCTION .......................................................................................................1 CHAPTER 1 .................................................................................................................3 INTRODUCTION ..............................................................................................................3 Overview .......................................................................................................5 Identification .................................................................................................8 Distribution ..................................................................................................16 Life History .................................................................................................20 Conservation ................................................................................................23 FIGURES ...............................................................................................................26 TABLES .................................................................................................................31 REFERENCES ........................................................................................................32 CHAPTER 2 ...............................................................................................................35 INTRODUCTION ....................................................................................................35 METHODS .............................................................................................................41 Morphological traits ....................................................................................41 DNA extraction, PCR amplification, and sequencing of mtDNA ..............41 Environmental Data .....................................................................................43 Blood properties ..........................................................................................44 Cloning and preliminary sequencing of globin genes .................................44 DNA extraction, PCR amplification, and sequencing of globins ................45 Characterization of hemoglobin isoform composition ................................46 ! vi! Measurement of hemoglobin functional properties .....................................47 RESULTS ...............................................................................................................48 Morphological traits ....................................................................................48 Population Genetic Analysis .......................................................................50 Molecular Dating and Population Delimitation ..........................................51 Environmental Data .....................................................................................54 Blood Properties ..........................................................................................56 Altitudinal patterns of sequence variation ...................................................58 Hemoglobin isoform composition ...............................................................59 Oxygenation properties of HbA and HbD isoforms ....................................60 Comparison of high- and low-altitude HbA and HbD variants ...................61 DISCUSSION ..........................................................................................................61 FIGURES ...............................................................................................................70 TABLES .................................................................................................................83 REFERENCES ........................................................................................................86 APPENDICES .........................................................................................................97 Appendix A .................................................................................................97 Appendix B ................................................................................................100
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