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Causes and Consequences of Sex Ratio Bias in Nazca

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Causes and Consequences of Sex Ratio Bias in Nazca CAUSES AND CONSEQUENCES OF SEX RATIO BIAS IN NAZCA BOOBIES (Sula granti) BY TERRI J. MANESS A Dissertation Submitted to the Graduate Faculty of WAKE FOREST UNIVERSITY GRADUATE SCHOOL OF ARTS AND SCIENCES in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY in the Department of Biology December 2008 Winston-Salem, North Carolina Approved By: David J. Anderson, Ph.D., Advisor ____________________________________ Examining Committee: Mark R. Leary, Ph.D., Chairman ____________________________________ Robert A. Browne, Ph.D ____________________________________ William E. Conner, Ph.D. ____________________________________ Clifford W. Zeyl, Ph.D. ____________________________________ ACKNOWLEDGEMENTS I thank the Galápagos National Park Service for permission to work in the Park and the Charles Darwin Research Station and TAME airlines for logistical support. For funding support, I am grateful to the Mead Foundation, the National Geographic Society, the National Science Foundation, the Oak Foundation, Sigma Xi, and Wake Forest University. I appreciate the work of my many colleagues for their work in producing the long-term Nazca booby databases, particularly Jill Awkerman, Tiffany Beachy, Julius Brennecke, Sebastian Cruz, Diego García, Kate Huyvaert, Elaine Porter, Devin Taylor, and Mark Westbrock. I also thank Julie Campbell and Amber Jones, who assisted with data entry and metabolite analyses, Matthew Furst, who also assisted with metabolite analyses, and Andrew D’Epagnier, who assisted with DNA extractions. I am particularly indebted to Audrey Calkins for her unmatched accuracy and speed of data entry, and for her endurance, skill, innovation, and unbounded zeal for PCR. Victor Apanius magnanimously shared his molecular analysis skills and vast statistical knowledge with me. I thank my lab mates, Jill Awkerman, Felipe Estela, Jacqueline Grace, Martina Müller, Elaine Porter, and Mark Westbrock for generously sharing their ideas, enthusiasm, knowledge, and friendship with me. I thank Drs. Robert Browne, William Conner, Mark Leary, and Clifford Zeyl for their encouragement and guidance. I am most grateful for the opportunity to work with Dr. David Anderson, an outstanding mentor and role model; I am a better biologist because of his encouragement, guidance, and support. CAUSES AND CONSEQUENCES OF SEX RATIO BIAS IN NAZCA BOOBIES (Sula granti) INTRODUCTION………………………………………………………………………...1 Nazca booby sex ratios............................................................................................1 Divorce rates in Nazca boobies...............................................................................4 Nazca booby mate choice........................................................................................6 Overview................................................................................................................11 Literature cited.......................................................................................................12 CHAPTER 1. ONTOGENIC SEX RATIO VARIATION IN NAZCA BOOBIES ENDS IN MALE-BIASED ADULT SEX RATIO.......................................................................18 Abstract........................................................................................................................19 Introduction.................................................................................................................20 Methods and Study Site..............................................................................................23 Statistical analyses.................................................................................................25 Results..........................................................................................................................27 Molecular sexing....................................................................................................27 Nestling survival....................................................................................................27 Return sex ratio......................................................................................................28 Discussion....................................................................................................................29 Acknowledgements.....................................................................................................31 Literature Cited..........................................................................................................32 ii Tables...........................................................................................................................36 Figures..........................................................................................................................37 CHAPTER 2. SERIAL MONOGAMY AND SEX RATIO BIAS IN NAZCA BOOBIES..........................................................................................................................40 Abstract........................................................................................................................41 Introduction.................................................................................................................42 Material and Methods................................................................................................46 Variance and skew in RS.......................................................................................46 Breeding state transition probabilities...................................................................47 Breeding bout lengths............................................................................................48 RS of divorced and retained males........................................................................48 Statistical analyses.................................................................................................49 Results..........................................................................................................................50 Variance and skew in RS.......................................................................................50 Breeding state transition probabilities...................................................................50 Breeding bout lengths............................................................................................51 RS of divorced and retained males........................................................................52 Discussion....................................................................................................................52 Acknowledgements.....................................................................................................57 References...................................................................................................................57 Tables...........................................................................................................................63 Figures..........................................................................................................................64 iii Appendix......................................................................................................................69 Figures....................................................................................................................69 Tables.....................................................................................................................73 CHAPTER 3. MATE ROTATION BY FEMALE CHOICE AND COERCIVE DIVORCE IN NAZCA BOOBIES (Sula granti)..............................................................75 Abstract........................................................................................................................76 Introduction.................................................................................................................77 Methods........................................................................................................................81 Behavioral observations.........................................................................................81 Reproductive success of male and female categories............................................87 Results..........................................................................................................................88 Mate rotation predictions.......................................................................................88 Male time out predictions......................................................................................93 Forced divorce predictions.....................................................................................93 Discussion....................................................................................................................95 Acknowledgements...................................................................................................100 References..................................................................................................................101 Tables.........................................................................................................................106 Figures........................................................................................................................108 Appendix....................................................................................................................112 iv CHAPTER 4. HEMATOLOGICAL AND MORPHOLOGICAL
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