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Dantzker Final Dissertation UNIVERSITY OF CALIFORNIA, SAN DIEGO Acoustic communication in the Greater Sage-Grouse (Centrocercus urophasianus) an examination into vocal sacs, sound propagation, and signal directionality A dissertation submitted in partial satisfaction of the requirements for the Doctor of Philosophy degree in Biology by Marc Steven Dantzker Committee in charge: Professor Jack W. Bradbury, Chair Professor Lin Chao Professor Grant Deane Professor James Moore Professor Sandra Vehrencamp 2015 © Marc Steven Dantzker, 2015 All rights reserved. The Dissertation of Marc Steven Dantzker is approved, and it is acceptable in quality and form for publication on microfilm and electronically: __________________________________________________________ __________________________________________________________ __________________________________________________________ __________________________________________________________ Chair University of California, San Diego 2015 iii DEDICATION For Heather, whose patience is nearly as unwavering as her love. For my father, mother, and sister, whose support I have too often required. For my kids, Charlie & Alice, who will henceforth address me as “Dr.” iv EPIGRAPH Before you cross the street take my hand. Life is what happens to you while you’re busy making other plans. -John Lennon v TABLE OF CONTENTS Signature Page ....................................................................................................... iii Dedication ............................................................................................................... iv Epigraph ................................................................................................................. v Table of Contents ................................................................................................... vi List of Figures ......................................................................................................... viii List of Tables .......................................................................................................... x Preface .................................................................................................................... xi Acknowledgements ................................................................................................ xvi Vita ......................................................................................................................... xviii Abstract of the Dissertation .................................................................................... xx Chapter 1: Vocal sacs and their role in avian acoustic display .............................. 1 Abstract ............................................................................................................. 1 Introduction ...................................................................................................... 2 Birds With Vocal Sacs ...................................................................................... 13 Vocal Sac Functions .......................................................................................... 35 References Cited ................................................................................................ 43 Chapter 2: Environmental Acoustics of Near Ground Communication: Acoustic Adaptation and the Greater Sage-Grouse, Centrocercus urophasianus ........... 50 Abstract ............................................................................................................. 50 Introduction ...................................................................................................... 51 Materials and Methods ...................................................................................... 54 vi Results ................................................................................................................ 64 Discussion .......................................................................................................... 73 Appendix ............................................................................................................ 87 List of Symbols .................................................................................................. 90 References Cited ................................................................................................ 92 Chapter 3: Directional Acoustic Radiation in the Strut Display Of Male Sage Grouse Centrocercus Urophasianus .............................................................................. 99 Summary ........................................................................................................... 100 Introduction ...................................................................................................... 100 Materials and Methods ...................................................................................... 101 Results ................................................................................................................ 105 Discussion .......................................................................................................... 108 Appendix A ........................................................................................................ 112 Appendix B ........................................................................................................ 113 Appendix C ........................................................................................................ 114 List of Symbols .................................................................................................. 114 References ......................................................................................................... 115 vii LIST OF FIGURES Chapter 1 Figure 1. Four types of tracheal elongation. ................................................................. 5 Figure 2. Tungara frog, Engystomops pustulosus, with inflated vocal sac. .................. 9 Figure 3. Laryngeal air sacs in primates. ....................................................................... 12 Figure 4. Esophageal vocal sac of the Rock Pigeon is inflated in the cooing courtship display. ..................................................................................................................... 15 Figure 5. Photographs of a dead American Bittern inflated and deflated. ................... 17 Figure 6. Bustard vocal sacs. ........................................................................................ 19 Figure 7. Vocal sacs of Umbrella birds. ....................................................................... 21 Figure 8. Grouse vocal sacs. ......................................................................................... 23 Figure 9. Ruddy Duck (Oxyura jamaicensis) thumping its air sac. .............................. 26 Figure 10. The air sacs of storks. ................................................................................... 28 Figure 11. The vocal sac of the Magnificent Frigatebird (Fregata magnificens). ........ 30 Figure 12. Gular pouch of the Pink-backed Pelican (Pelecanus rufescens). ................. 32 Figure 13. Illustration of the booming display of the Kakapo (Strigops habroptilus). 34 Chapter 2 Figure 1. Source-receiver geometry for an acoustic sender and receiver. ..................... 56 Figure 2. Comparisons of model predictions with measured transmission loss. ........... 66 Figure 3. Transmission Loss as a function of time and temperature. ........................... 69 Figure 4. Varying the effective flow resistivity of the ground. .................................... 72 viii Figure 5. Model predictions of how acoustic transmission changes as a female approaches a male. ................................................................................................... 77 Figure 6. Model predictions of how acoustic transmission changes with different common female and male behaviors. ........................................................................ 81 Chapter 3 Figure 1. Strut display of male sage grouse. .................................................................. 101 Figure 2. Spectrogram of the acoustic component of a male sage grouse strut. ........... 101 Figure 3. Schematic drawing of the acoustic array deployment ................................... 103 Figure 4. Transmission loss functions for 12 measurements. ....................................... 104 Figure 5. Transmission loss curves for all propagation paths. ...................................... 106 Figure 6. Acoustic beam patterns for the coos and pops. ............................................. 108 Figure 7. Polar histograms showing the orientation of the beam patterns’ major features for coo 1 and coo 2. .. ................................................................................................ 109 Figure 8. Time-varying beam patterns for the whistle component. .............................. 110 Figure 9. Polar histograms showing the orientation of the beam patterns’ major features for the whistle... ........................................................................................................ 111 ix LIST OF TABLES Chapter 2 Table 1. Ranges of effective flow resistivity, σe , for various types of ground surface according to Embleton et al., 1983 ........................................................................... 67 Chapter 3 Table 1. Directionality of the coo and pop notes of the sage grouse strut display. ...... 107 Table 2. Results of Rayleigh tests
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