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The University of Chicago Evolution and Sociality In THE UNIVERSITY OF CHICAGO EVOLUTION AND SOCIALITY IN FAIRY-WRENS (AVES: MALURIDAE) A DISSERTATION SUBMITTED TO THE FACULTY OF THE DIVISION OF THE BIOLOGICAL SCIENCES AND THE PRITZKER SCHOOL OF MEDICINE IN CANDIDACY FOR THE DEGREE OF DOCTOR OF PHILOSOPHY DEPARTMENT OF ECOLOGY AND EVOLUTION BY ALLISON ELAINE JOHNSON CHICAGO, ILLINOIS AUGUST 2016 TABLE OF CONTENTS LIST OF TABLES ............................................................................................................. iv LIST OF FIGURES ........................................................................................................... vi ACKNOWLEDGMENTS ................................................................................................ vii CHAPTER 1: EVOLUTION AND SOCIALITY IN FAIRY-WRENS (AVES: MALURIDAE) ............................................................................................................................................. 1 INTRODUCTION ................................................................................................................. 1 CONCLUSIONS ................................................................................................................... 6 CHAPTER 2: DIFFERENT MODES OF EVOLUTION IN MALES AND FEMALES GENERATE DICHROMATISM IN FAIRY-WRENS (MALURIDAE) .......................... 9 ABSTRACT ......................................................................................................................... 9 INTRODUCTION ............................................................................................................... 10 METHODS ........................................................................................................................ 12 RESULTS ......................................................................................................................... 19 DISCUSSION ..................................................................................................................... 27 APPENDIX 2A: PLUMAGE SCORES FOR MALURIDAE SPECIES ............................................. 34 CHAPTER 3: HETEROSPECIFIC SOCIALITY MEDIATED BY SONG DISCRIMINATION IN FAIRY-WRENS .......................................................................................................... 44 ABSTRACT ....................................................................................................................... 44 INTRODUCTION ............................................................................................................... 45 METHODS ........................................................................................................................ 47 ii RESULTS ......................................................................................................................... 54 DISCUSSION ..................................................................................................................... 62 APPENDIX 3A: LOADINGS DERIVED FROM PCA ON RESPONSE VARIABLES FOR HETEROSPECIFIC AND CONSPECIFIC RECOGNITION EXPERIMENTS ...................................................................... 66 APPENDIX 3B: CROSS TABULATION OF SPLENDID AND VARIEGATED NESTING SUCCESS AND RE- NESTING BEHAVIOR ON SHARED AND SOLITARY TERRITORIES ......................................... 67 APPENDIX 3C: RAW RESPONSES OF SPLENDID AND VARIEGATED FAIRY-WRENS TO HETEROSPECIFIC AND CONSPECIFIC PLAYBACK EXPERIMENTS ................................................................... 68 CHAPTER 4: HELPING BEHAVIOR AND PROMISCUITY IN THE VARIEGATED FAIRY- WREN: SEX SPECIFIC EFFECTS OF AUXILIARY MEMBERS ............................... 72 ABSTRACT ....................................................................................................................... 72 INTRODUCTION ............................................................................................................... 73 METHODS ........................................................................................................................ 76 RESULTS ......................................................................................................................... 84 DISCUSSION ..................................................................................................................... 91 LITERATURE CITED ..................................................................................................... 97 iii LIST OF TABLES Table 2.1. Summary of outputs from evolutionary models of plumage evolution ........... 24 Table 2.A.1. Plumage scores for males of Maluridae species .......................................... 34 Table 2.A.2. Plumage scores for females of Maluridae species ....................................... 39 Table 3.1. Time allocation for splendid and variegated fairy-wrens when along and when with co-resident ......................................................................................................................... 56 Table 3.2. Pairwise comparisons of response to heterospecific playback stimuli ............ 60 Table 3.3. Pairwise comparisons of response to conspecific playback stimuli ................ 61 Table 3.A.1. Principle components analysis loadings of response variables to heterospecific stimuli ............................................................................................................................... 66 Table 3.A.2. Principle components analysis loadings of response variables to conspecific stimuli ........................................................................................................................................... 66 Table 3.B.1. Cross tabulation fledging and re-nesting behavior for birds in shared and solitary territories ........................................................................................................................... 67 Table 3.C.1. Raw responses of splendid fairy-wrens to heterospecific song stimuli ....... 68 Table 3.C.2. Raw responses of variegated fairy-wrens to heterospecific song stimuli .... 69 Table 3.C.3. Raw responses of splendid fairy-wrens to conspecific song stimuli ............ 70 Table 3.C.4. Raw responses of variegated fairy-wrens to conspecific song stimuli ........ 71 Table 4.1. Microsatellite loci used in paternity analyses .................................................. 80 Table 4.2. Variability and exclusion probabilities of microsatellite loci .......................... 81 iv Table 4.3. Nominal logistic models of fledging success and extra-pair paternity ............ 84 Table 4.4. Standard least squares models of egg and nestling size and feeding rate ........ 87 Table 4.5. Extra-pair paternity rates and proportion of extra-pair young across years .... 90 v LIST OF FIGURES Figure 2.1. Phylogeny depicting evolutionary changes in plumage characters ................ 21 Figure 2.2. Pair-wise plumage distances between taxa as a function of divergence ........ 22 Figure 2.3. Ancestral reconstruction of geographic state ................................................. 25 Figure 2.4. Pair-wise comparisons between dichromatism and breeding latitude ............ 26 Figure 3.1. Percentage of splendid and variegated fairy-wren groups fledging young from their first nest ............................................................................................................................. 55 Figure 3.2. Frequency of vigilance of splendid and variegated fairy-wrens when in the presence and absence of co-resident ................................................................................................ 57 Figure 3.3. Strength of response to heterospecific playback stimuli ................................ 59 Figure 3.4. Example response to heterospecific song stimuli ........................................... 60 Figure 3.5. Strength of response to conspecific playback stimuli .................................... 61 Figure 4.1. Fledging success as a function of group size .................................................. 85 Figure 4.2. Mean egg volume as a function of male or female auxiliaries ....................... 86 Figure 4.3. Dominant pair feeding rate as a function of male or female auxiliaries ........ 89 Figure 4.4. Extra-pair paternity rate as a function of the number of male auxiliaries ...... 91 vi ACKNOWLEDGMENTS My dissertation was made possible by many people who helped and supported me during my tenure at the University of Chicago. I am extremely indebted to my advisor, Stephen Pruett- Jones, who has helped me at every step of the way. My committee, Jill Mateo, Trevor Price, John Bates, and Tim Wootton have helped in the development, writing, and analysis of this work. Christina Masco helped generate song stimuli, design experiments, teach recording techniques, and was a wonderful collaborator and confidant in the field. Niles Desmarais, Brian Gallagher, Emily Geary, Ian Hoppe, Sara Jackrel, Eva Kinnebrew, Dylan Meyer, Jenny Uehling, Kelle Urban, and Peter Zaykoski assisted in many hours of field-work (with special thanks to Dylan who came to Brookfield to keep me sane three seasons in a row). Terry Peacock and Tricia Curtis, as well as the rest of the Conservation Volunteers Australia family made access to Brookfield Conservation Park both possible and a joy, and protected us from the heat and the bushfires. Jill Mateo loaned essential equipment for the fieldwork and assisted with experimental design. Jordan Price helped design and carry out analyses for Chapter 1. Jon Mitchell provided statistical and coding support.
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