Ecology of Wintering Black-Capped Vireos in Mexico
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ECOLOGY OF WINTERING BLACK-CAPPED VIREOS IN MEXICO A Dissertation by ROBERT ANDREW POWELL Submitted to the Office of Graduate and Professional Studies of Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Chair of Committee, R. Douglas Slack Committee Members, William E. Grant R. Neal Wilkins X. Ben Wu Head of Department, Michael Masser December 2013 Major Subject: Wildlife and Fisheries Sciences Copyright 2013 Robert Andrew Powell ABSTRACT The black-capped vireo (Vireo atricapilla) is an endangered Neotropical migratory songbird that has received considerable attention in its breeding range, but relatively little attention in its winter range in Mexico. To address information needs regarding winter ecology of the black-capped vireo, I conducted research in Mexico focused on investigations of the winter distribution, habitat use, and migratory linkages between breeding and wintering sites. Over 3 winter periods in 2002–2004, I identified and described the geographic distribution for the black-capped vireo at study sites across 8 states in western Mexico and determined if differential migration occurs among different classes of individuals. I documented winter occupancy in the 5 most northern Mexican states surveyed (Sinaloa, Durango, Nayarit, Jalisco, and Colima), and identified a strong association between sex/age class and winter latitude with adult males occupying habitat at more northern latitudes relative to females and juveniles. During two winters in 2003-2004, I conducted field research at study sites in Mexico to evaluate patterns of winter habitat use by black-capped vireos and determine which habitat characteristics may influence vireo use of winter patches. Winter habitat use by black-capped vireos was best predicted by increasing values of slope and foliage cover, and by decreasing values of canopy cover and tree diameter. Vireo use plots characterized as thorn forest had greater foliage density, greater shrub density, less canopy cover, and smaller tree diameter than plots classified as tropical deciduous or ii semi-deciduous forest, suggesting that thorn forests may be most suitable for vireo occupancy during the winter months. I also used stable carbon, hydrogen, and nitrogen isotopes in black-capped vireo feathers obtained across 3 states in the breeding range and 8 states in the wintering range to determine if vireo feathers collected on the wintering grounds could be used to assess breeding origins and to determine if a relationship exists between breeding and wintering latitudes such as chain or leapfrog migration. Feathers collected at both breeding and wintering sites displayed considerable variability in isotopic composition for all 3 isotopes analyzed, and thus did not provide sufficient information to establish migratory linkages between breeding and wintering sites. iii DEDICATION To my family Mom For your unwavering love and support in all that I do Dad For wanting this for me, I wish you could still be here Marilyn For the happiness you bring to me iv ACKNOWLEDGEMENTS I sincerely appreciate the generous financial support for this research provided by the U.S. Fish and Wildlife Service (USFWS) and Texas Parks and Wildlife Department (TPWD). I especially thank Dr. Craig Farquhar of TPWD who first envisioned a study of vireos in Mexico and who was instrumental in many ways in getting this project off the ground. Thanks as well to the Departments of Wildlife and Fisheries Sciences (WFSC), Biology, and Rangeland Ecology and Management (now Ecosystem Science and Management; ESSM), as well as the Texas Wildlife and Fisheries Sciences Extension Unit who all have employed me at various times throughout my time at Texas A&M University (TAMU). I also wish to acknowledge the USFWS Migratory Bird Permit Office, the USFWS Division of Management Authority, and the Secretaría de Medio Ambiente y Recursos Naturales (SEMARNAT) for permits, as well as Dr. Felipe Chavez-Ramirez for granting me use of his bird banding permit. I am indebted to Dr. José Ignacio “Nacho” González and Oscar Ballesteros at the Universidad Autónoma de Nuevo León for storing feathers for me, obtaining expert permits, and for assistance with preparing and exporting the feathers across the border. To Dr. Xico Vega of PRONATURA and the Universidad Autónoma de Sinaloa, I appreciate the access they provided to La Reserva Ecológica del Mineral de Nuestra Señora in Cosalá, Sinaloa. I extend my gratitude as well to Marcela Ley from PRONATURA and her team of young biologists who accompanied us to the Reserve, v provided campsites, food, and entertainment. My gratitude extends as well to Dr. Eduardo Santana Castellón for access to la Reserva de la Biosfera Sierra de Manantlán. I am grateful to the generous research museums and collections that provided feather tissues for isotopic analysis. The staff and museums to which I owe my thanks include Dr. Carla Cicero at the University of California, Berkeley Museum of Vertebrate Zoology, Mark Blair Robbins at the University of Kansas Natural History Museum, Paul Sweet at the American Museum of Natural History, Dr. David Willard at the Field Museum of Natural History, Gene Hess at the Delaware Museum of Natural History, Dr. John Hafner at the Moore Laboratory of Zoology at Occidental College, Dr. Robert Prys-Jones at the British Museum of Natural History, and Dr. Gary Graves at the Smithsonian Institution National Museum of Natural History. I also owe tremendous gratitude to Dr. David Cimprich of The Nature Conservancy and Dr. Joseph Grzybowski of the University of Central Oklahoma, who collected and provided feather samples from Fort Hood, Texas, and Fort Sill, Oklahoma, respectively. The Colorado Plateau Stable Isotope Laboratory (CPSIL) in Flagstaff, Arizona conducted the feather isotope analyses for me, and I am indebted to Richard Doucett of CPSIL for his guidance and cooperation. I appreciate the cooperation of Leslie Schwierzke Wade, Reagan Errera, and Carrie Miller in Dr. Dan Roelke’s laboratory in WFSC at TAMU for access to space and equipment used for preparation of feather tissues for isotopic analysis. I am forever grateful to my field assistants Chris Merkord, Jennifer Reidy, and Chris Burney, who accompanied me to Mexico and each contributed tremendously to the vi success of the project. Others who were helpful in various aspects of this research include Dr. Keith Hobson with Environment Canada, who generously provided his time for conversations about my isotope research, particularly in helping trying to understand some of the problems I encountered with my results. Dr. Miguel Mora of WFSC also provided useful feedback on the isotope data and other aspects of my research. To the many bird enthusiasts on listservs who provided tips as to where to look for black- capped vireos in Mexico, I owe my sincere gratitude. Also, I greatly appreciate the generosity of the expatriates in Mexico who were interested in my project, particularly Ben Wassink who was a local guru on the birds around Laguna Santa Maria del Oro in Nayarit. He and others at Koala Bungalows often provided us food and good company as well. To the people of Mexico, I cannot express enough how much I admire and appreciate your generosity and goodwill. I would like to thank the members of my committee for their guidance and support throughout the course of my research, including Dr. Douglas Slack (WFSC), Dr. Bill Grant (WFSC), Dr. Neal Wilkins (WFSC), and Dr. Ben Wu (ESSM). I particularly want to express my gratitude towards Dr. Slack, who in addition to being my Committee Chair, I also consider a friend, colleague, and mentor. The education he provided me extends far beyond this dissertation or any coursework. Staff of WFSC at TAMU helped keep me on track throughout my studies; I thank all of them, especially Vicki Buckbee, Shirley Konecny, and Diann Radke. Thanks also to my many friends and colleagues at TAMU who have made my time here a terrific experience. I owe a debt of thanks to each of my lab mates in the vii Slack lab for their friendship, humor, advice, and much more, including April Conkey, Danielle Rutka Greer, Dan Kim, Kristin Millenbach LaFever, Glenn Proudfoot, Dawn Sherry, Tsuyoshi Watanabe, and Laura Weber. I also wish to thank Glenn for his tremendous assistance with preparing and building various pieces of equipment for my fieldwork in Mexico. I appreciate the assistance provided by Heather Mathewson, Paige Schmidt, and Tiffany McFarland with other aspects of my research. The list of friends who have provided me friendship while at TAMU is lengthy, but in addition to my lab mates and others mentioned above, other friends I wish to thank include Kevin Baker, Lindsay Bennett, Meg Byerly, Thomas Dixon, Lance and Eve Fontaine, Richard Heilbrun, Christian Kaufman, David LaFever, Eddie Lyons, Anna Munoz, Dean Ransom, Christa Iacono Ruffins, Angela Skeeles-Worley, Stacey Allison Steinbach, and Holly Stephens. I especially thank my family for their unconditional love, support, patience, and encouragement. Thank you to my mother, Mary Craig Powell, my brother John, his wife Wendy, and their daughter Rebecca. Thank you to my dogs Maggie (now deceased) and Scout, who were always by my side. Finally, thank you to Marilyn Stone for your love and encouragement; you truly are special to me. viii NOMENCLATURE ‰ Per mil 13C Stable carbon isotope 2H Stable hydrogen isotope (deuterium) 15N Stable nitrogen isotope AICc Akaike’s Information Criterion for small sample sizes ANOVA