Transmission of Structurally Distinct Song Phrases in the White-Crowned Sparrow (Zonotrichia leucophrys pugetensis) Thesis Presented in Partial Fulfillment of the Requirements for the Degree Master of Science in the Graduate School of The Ohio State University By Erica Marie Szeyller-Macolley, B.A. Graduate Program in Evolution, Ecology, and Organismal Biology The Ohio State University 2012 Master’s Examination Committee: Dr. Douglas A. Nelson, Advisor Dr. W. Mitch Masters Dr. J. Andrew Roberts Dr. Angelika Poesel Copyright by Erica Marie Szeyller-Macolley 2012 Abstract Birdsong in many species has evolved to transmit over long distances to potential mates and competitors. Song can be structurally complex, often consisting of many different components. Variation in the frequency and duration of these components may influence attenuation and degradation of songs as they transmit through the environment. Habitat structure will also affect attenuation and degradation of certain song traits and influence what information is available for receivers. To understand how variation in song and habitat structure may affect song degradation during transmission, we broadcast songs of the Puget Sound white-crowned sparrow, Zonotrichia leucophrys pugetensis, in two different habitats commonly occupied by the species. The song of this species contains four structurally and functionally distinct phrases (whistle, note complex, buzz, and trill), thirteen examples of which were broadcast over two biologically relevant distances (48 m and 96 m). Spectrogram cross-correlation between the song phrases recorded at 48 and 96 m, and undegraded model songs provided a measure of song degradation. Overall, sound attenuation and degradation increased from 48 to 96 meters. Transmission fidelity differed among phrases; in particular, whistle phrases showed less degradation and attenuation than other phrases. Despite the high average transmission fidelity, whistle phrases were misclassified more often and were most similar among ii phrase examples than any other phrase, indicating that simple whistles may be potentially difficult for receivers to discriminate amongst. Conversely, the acoustically diverse note complex phrases were the least likely to be misclassified and were most dissimilar among phrase examples. Transmission of song phrases differed between the two habitats. Phrases were more attenuated and degraded as well as more similar after degradation in an open dune habitat (Bullard’s Beach) while there was more reverberation in a forest edge habitat (Cape Blanco). Lastly, phrases attenuated and degraded similarly within each site. Certain phrases propagate and are potentially perceived differently than others, thus supporting a correlation between signal structure and function. Simple tonal whistles resist attenuation and degradation and may alert listeners to the individual- and dialect- identifying information in acoustically complex note complexes and trills that follow. While the note complex and trill phrase attenuate and degrade more than the whistle phrase over transmission, their structural complexity nevertheless may contain sufficient redundancy that allows them to be more easily discriminated. iii Acknowledgements First and foremost, I am thankful for the guidance and support of my advisor, Douglas A. Nelson as well as for the thoughtful advice from present and past committee members, Angelika Poesel, W. Mitch Masters, J. Andrew Roberts, and Amanda Rodewald. A number of graduate students helped immensely with my project. In particular, I would like to thank Jimmy Chiucchi, Brandon Sinn, and Jessica Hall for their statistical, writing, and presentation guidance. Also, thanks to Stephanie Wright, Dee Bolen, Desiree Narango, and Laura Kearns for lively pizza-fueled discussions on theory and methods in animal behavior. At Ohio State, I have been fortunate to work with a number of inspiring teachers that have helped me learn and grow as an instructor. In particular, I want to thank Cindy Bronson, Jimmy Chiucchi, John Harder, Douglas A. Nelson, Angelika Poesel, Joe Raczkowski, and Judy Ridgway. Throughout my time here, I have been lucky enough to surround myself with friends & family who have provided me with unwavering emotional support, confidence, and lots of laughter through this process. Most special thanks, to my husband Brent for seeing me through every step of this process with patience, thought, understanding, and most importantly humor. iv Vita June 2004 .......................................................B. Reed Henderson High School May 2008 .......................................................B.A. Psychology, West Chester University August 2008 to present .................................Graduate Teaching Associate, Department of EEOB, The Ohio State University Awards 2010-2011 Dept. of Evolution, Ecology, and Organismal Biology Graduate Teaching Award. Presented June 2011. Publications Published Abstracts of Presentations at Professional Meetings Johnson, V. K., Gans, S. E., Kerr, S., LaValle, W., Bee, S., Szeyller, E., and Wonsock, G. (March, 2007). Emotion coping and adolescent behavior problems: Validating the TMMS. Poster presented at the Biennial Meeting of the Society for Research in Child Development, Boston, MA. Szeyller-Macolley, E., Poesel, A., and Nelson, D. A. (August, 2011). Transmission of structurally distinct song phrases in the white-crowned sparrow. Poster presented at the Joint Meeting of the Animal Behavior Society and the International Ethological Conference, Bloomington, IN. Szeyller-Macolley, E., Poesel, A., and Nelson, D. A. (April, 2012). Transmission and reception of structurally distinct song phrases in the white-crowned sparrow (Zonotrichia leucophrys pugetensis). Talk presented at the 19th Annual Animal Behavior Conference, Bloomington, IN. v Fields of Study Major Field: Evolution, Ecology, and Organismal Biology vi Table of Contents Abstract ............................................................................................................................... ii Acknowledgements ............................................................................................................ iv Vita ...................................................................................................................................... v List of Tables ..................................................................................................................... ix List of Figures ..................................................................................................................... x Chapter 1: Acoustic Communication in Songbirds ............................................................ 1 Introduction ..................................................................................................................... 1 Signaling Modalities ....................................................................................................... 2 Properties of Sound ......................................................................................................... 5 Song Production .............................................................................................................. 6 Sound Transmission ........................................................................................................ 7 Sound Reception ........................................................................................................... 11 Signaling Ecology ......................................................................................................... 13 Song Structure Function and Evolution ........................................................................ 14 Conclusion ..................................................................................................................... 17 Chapter 2: Transmission of Structurally Distinct Song Phrases in the White-Crowned Sparrow (Zonotrichia leucophrys pugetensis) .................................................................. 19 Introduction ................................................................................................................... 19 Methods ......................................................................................................................... 23 Sites ........................................................................................................................... 23 Sound File Preparation .............................................................................................. 23 Sound Transmission Trials ........................................................................................ 24 Sound Analysis .......................................................................................................... 25 Results ........................................................................................................................... 27 Interaction of Sites and Song Phrases........................................................................ 27 Song Phrases .............................................................................................................. 28 vii Sites ........................................................................................................................... 30 Discussion ..................................................................................................................... 32 Interaction of Sites and Song Phrases.......................................................................