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Phylogenetic and ecological aspects of cooperative breeding in the bee-eaters (Aves: Meropidae) Item Type text; Dissertation-Reproduction (electronic) Authors Burt, Donald Brent, 1965- Publisher The University of Arizona. Rights Copyright © is held by the author. Digital access to this material is made possible by the University Libraries, University of Arizona. Further transmission, reproduction or presentation (such as public display or performance) of protected items is prohibited except with permission of the author. Download date 27/09/2021 02:11:00 Link to Item http://hdl.handle.net/10150/282167 INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quali^ of this reproduction is dependent upon the quality of the copy submitted. 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Contact UMI directly to order. UMI A Bell & Howell Information Company 300 North Zedj Road, Ann Aibor MI 48106-1346 USA 313^61-4700 800/521-0600 PHYLOGENETTC AND ECOLOGICAL ASPECTS OF COOPERATIVE BREEDING IN THE BEE-EATERS (AVES: MEROPIDAE) by Donald Brent Bxirt A Dissertation Submitted to the Faculty of the DEPARTMENT OF ECOLOGY AND EVOLUTIONARY BIOLOGY In Partial Fvdfillment of the Requirements For the Degree of DOCTOR OF PHILOSOPHY In the Graduate College THE UNIVERSITY OF ARIZONA 1996 DMI Nxjinber; 9713401 UMI Microform 9713401 Copyright 1997, by UMI Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. UMI 300 North Zeeb Road Ann Arbor, MI 48103 2 THE UNIVERSITY OF ARIZONA GRADUATE COLLEGE As members of the Final Examination Committee, we certify that we have read the dissertation prepared by tVinalH Rr-^nt-. Rnr-h entitled Phyloqenetic and Ecological Aspects; of O^ioperative Breeding in the Bee-eaters (Aves: Meropida^K and recommend that it be accepted as fulfilling the dissertation requirement for the Degree of of Philosophy Date ' ~ i!.. 7/zz/9e Date Date Final approval and acceptance of this dissertation is contingent upon the candidate's submission of the final copy of the dissertation to the Graduate College. I hereby certify that I have read this dissertation prepared under my direction and recommend that it be accepted as fulfilling the dissertation requirement. DissertaG^n Director Date ' Wayne I-Iaddison 3 STATEMENT BY AUTHOR This dissertation has been submitted in partial fulfillment of requirements for an advanced degree at The University of Arizona and is deposited in the University Library to be made available to borrowers under rules of the Library. Brief quotations from this dissertation are allowable without special permission, provided that accurate acknowledgment of source is made. Requests for permission for extended quotation from or reproduction of this manuscript in whole or in part may be granted by the head of the major department or the Dean of the Graduate College when in his or her judgment the proposed use of the material is in the interests of scholarship. In all other instances, however, permission must be obtained from the author. SIGNED: / 4 ACKNOWLEDGMENTS I thank my committee members: Judie Broiistein, David Maddison, Nancy Moran, Dan Papaj, and the chair, Wayne Maddison, for their advise and close attention to my dissertation research. I received considerable advise on bee-eater biology, behavioral phylogenetics, and avian systematica from people outside the University of Arizona including Alan de Queiroz, Stephen Emlen, Hilary Fry, Brad Livezey and Doug Siegel-Causey. Groups deserving special mention included the Phylogeny Discussion Group and the entire crew in Wayne Maddison's lab, especially Greta Binford, Marshal Hedin, Susan Masta and Shelley McMahan. Patrick O'Grady and Joana Silva were also always encouraging and supportive. Very special thanks go out to Wade Leitner for getting me out into the field so that I did not forget what birds look like without cotton eyes. Crucial therapy was provided by the departmental basketball and ultimate Frisbee groups and my many dancing partners, particularly Sarah Mesnick. Consistent and considerable financial support was provided by the Department of Ecology and Evolutionary Biology, the Silliman Memorial fund, and particularly the Research Training Group in the Analysis of Biological Diversification (Fellowships and research grants) and the National Science Foimdation, (Dissertation Improvement Award). Field Work in Kenya was made possible with the assistance of Terry Repp and Kim Score from the University of Arizona, and Titus Imboma, Peter Njoroge, Edward Wayaki and the remaining staff of National Museums of Kenya. In Thailand, Kim Burt worked very hard in the field. I was also assisted by Dr. Schwann of the Royal Forest Department, and Dr. Sawat, Mr. Som Yot, and Mr. Boriphan, of the National Watershed Management Division. Dr. Jamroon and Ms. Piyathip of the National Marine Parks Division were also very accommodating. The following Museums were gracious in either loaning specimens or allowing me access to their collections: the University of Kansas Museum of Natural History, the National Museum of Natural History, and the British Museimi of Natural History. Finally, this degree and the work behind it would not have been possible without the ever present support of my family. Kim Burt provided a great deal of support in so many ways for which I will always be appreciative. DEDICATION This work is dedicated to my anchors in life, the people who are always ready with their love and support; my family. 6 TABLE OF CONTENTS LIST OF HGURES 8 LIST OF TABLES 10 ABSTRACT 11 CHAPTER ONE- INTRODUCTION TO THE EVOLUTION AND MAINTENANCE OF COOPERATIVE BREEDING IN BIRDS 13 FOUR BY FOUR; LEVELS OF ANALYSIS AND MAJOR QUESTIONS 15 WHY DO HELPERS DELAY DISPERSAL? 26 Selective advantages 26 Proximate mec±ianisms 26 OntQggny 27 WHY DON'T HELPERS BREED INDEPENDENTLY? 28 Selective advantages, helper viewpoint 28 Potential conflict of interest with breeders 30 Proximate mechanisms 31 Ontogeny 31 WHY DO INDIVIDUALS HELP? 33 Selective advantages 33 Proximate mechanisms 34 Ontogeny 35 DO HELPERS REALLY HELP? 36 HISTORICAL PATTERNS 37 Phylogenetic constraints 38 SUMMARY 38 CHAPTER TWO- PHYLOGENETIC RELATIONSHIPS OF THE BEE-EATERS (AVES: MEROPIDAE) 40 METHODS 43 Plumage characters 43 Phvlogenetic analyses 46 RESULTS 51 DISCUSSION 59 Comparison of primary and secondary analyses 59 Biogeographic patterns 60 Comparison with earlier systematic study 64 Conclusions 69 CHAPTER THREE- BREEDING BIOLOGY AND SOCIALITY OF BEE-EATERS IN THAILAND 70 STUDY SITES AND METHODS 71 RESULTS 73 Little green bee-eaters. 73 Colony structure 73 Breeding chronology 73 Predators and anti-predatory behavior 73 7 TABLE OF CONTENTS - Continued Evidence of cooperative breeding. 74 Blue-tailed bee-eaters. 75 Colony structure 75 Breeding chronology 75 Evidence of cooperative breeding and sociality 75 Bay-headed and blue-bearded bee-eaters. 76 DISCUSSION 78 Little green bee-eaters in India and Thailand 78 Social systems of poorly studied spedes 79 Significance of these data in comparative studies 80 CHAPTER FOUR- BEHAVIORAL MALLEABILITY VERSUS PHYLOGENETIC INERTLA. AND COOPERATIVE BREEDING IN BEE- EATERS 83 METHODS 89 RESULTS 93 Patterns of evolution in breeding systems 93 Correspondence of other traits to breeding system evolution 99 Social systems 99 Nesting substrate requirements 102 Habitat utilization 104 Wing shape, migration, and foraging mode 106 Diet specialization 113 DISCUSSION 115 Evolution of breeding systems relative to other traits 115 Behavioral malleability versus phylogenetic inertia 119 Need for detailed ecological data on all species 122 Summary 124 CHAPTER FIVE- PHYLOGENETIC STRUCTURE INDICES: WHAT DO THEY TELL US AND HOW SHOULD THEY BE USED? 126 Expectations of a preferred index 128 METHODS 133 RESULTS 134 Matrices failure rate 134 Behavior of CI. RI. and RC 136 Behavior of gi 138 Behavior of average nonparametric bootstrap values 140 DISCUSSION 142 CONCLUSIONS 146 APPENDIX A 148 APPENDIX B 157 APPENDIX C 172 WORKS CITED 173 8 LIST OFnCURES FIGURE 2.1, Consensus tree from primary analyses of races 52 FIGURE 2.2, Four trees from primary analyses of species 53 FIGURE 2.3, Trees from primary an^yses with standardized character weights 54 FIGU^ 2.4, Trees from secondary analyses where character weights equaled one 56 FIGURE 2.5, Trees from successive approximation character weighting analyses 57