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Thesis 7Apr13 The Role of Male Vocal Signals During Male-Male Competition and Female Mate Choice in Greater Prairie-Chickens (Tympanuchus cupido) Thesis Presented in Partial Fulfillment of the Requirements for the Degree Master of Science in the Graduate School of The Ohio State University By Jennifer Ann Hale, B. S. Graduate Program in Evolution, Ecology, and Organismal Biology The Ohio State University 2013 Thesis Committee: Dr. Jacqueline Augustine, Advisor Dr. Douglas Nelson, Advisor Dr. Andrew Roberts Dr. William Mitchell Masters Copyright by Jennifer Ann Hale 2013 Abstract In many taxa, vocal communication plays an integral role in aggression, territorial defense, and female choice. The acoustic structure of vocalizations is influenced by physical constraints on the vocalizer, suggesting a potential for discrimination among individuals. In the lek-mating Greater Prairie-Chicken (Tympanuchus cupido), male vocalizations are an integral part of the ritualized display. We investigated whether variation among vocal characteristics of individual male Greater Prairie-Chickens plays a role during female choice and male-male competition. Vocal characteristics varied among males but were fairly consistent for each male, suggesting that vocalizations might be used by prairie-chickens to identify individuals. Female choice was evaluated by comparing characteristics of vocalizations produced by reproductively successful and unsuccessful males, and successful males were found to vocalize at a relatively lower mean frequency. Playbacks of familiar and unfamiliar males were conducted on the lek to assess the role of vocalizations during male-male competition. Males responded to the prairie-chicken treatments by vocalizing at a faster rate and approaching the playback speaker, but they did not respond more strongly to the vocalizations of unfamiliar males than familiar males. Our results suggest that variation is present ii among the vocalizations of individual male Greater Prairie-Chickens and that this variation could be used by females during mate choice, but signal variation does not appear to be used by males to discriminate among familiar individuals and strangers. However, vocalization elicits an aggressive response in males that hear it, regardless of the individual that has produced it. Vocalization likely functions as a way of announcing that a territory is occupied and defended, but it may also serve as a way of advertising to male or female conspecifics or as a signal that is secondary to other forms of communication. iii Acknowledgments I would like to thank my advisor, Dr. Jackie Augustine, for her invaluable assistance, mentorship, and guidance during all stages of this project. I would also like to thank my co-advisor, Dr. Doug Nelson, for his great help and feedback along the way. My other committee members, Dr. Andy Roberts and Dr. Mitch Masters, contributed a great deal of expertise that strengthened the project. I am grateful to Konza Prairie Biological Station, Rannell’s Ranch Preserve, and the private landowners who allowed me to conduct research on their properties. Funding in the form of an OSU-Lima Research and Special Projects Grant, an NAOC Student Travel Award, and travel money from the department of Evolution, Ecology, and Organismal Biology at The Ohio State University made it possible for me to conduct fieldwork in Kansas and travel to scientific conferences. Finally, I would like to thank my parents, my boyfriend Marc, and many friends and family members for their unfailing kindness, patience, and good humor. I am so very fortunate to have had their support over the years and cannot thank them enough. iv Vita 2010 ........................................................................... B.S. Biology, Arizona State University 2010-2013 ............... Graduate Teaching Assistant, Department of Evolution, Ecology, and Organismal Biology, The Ohio State University Fields of Study Major Field: Evolution, Ecology, and Organismal Biology v Table of Contents Abstract .............................................................................................................................. ii Acknowledgments ............................................................................................................. iv Vita ..................................................................................................................................... v List of Tables ..................................................................................................................... vii List of Figures ................................................................................................................... viii Introduction ........................................................................................................................ 1 Methods ............................................................................................................................. 5 Results .............................................................................................................................. 11 Discussion ......................................................................................................................... 17 Literature Cited ................................................................................................................. 23 Appendix A: Tables ........................................................................................................... 28 Appendix B: Figures .......................................................................................................... 36 vi List of Tables Table 1: Results of principal component analysis using vocal characteristics .................. 28 Table 2: Results of a linear mixed model analysis of the relationship between vocal characteristics of 56 male Greater Prairie-Chickens and date, year, and female presence with male ID as a random effect ................... 29 Table 3: Results of discriminant function analyses including 5 vocalizations from each of 45 males ........................................................................................... 30 Table 4: Results of principal component analysis using behavioral characteristics ........ 31 Table 5: Results of a linear mixed model analysis of the relationship between environmental factors and behavior for 74 males with male ID as a random effect .................................................................................. 32 Table 6: Coefficients of variation for vocal, behavioral, and morphological characteristics of male Greater Prairie-Chickens near Manhattan, Kansas in 2011 and 2012 .......................................................................................... 33 Table 7: Summary statistics for differences in response measures before and during treatments in 10 playback trials, each including 4 treatments: owner, neighbor, stranger, and control ........................................... 34 Table 8: Results of principal component analysis using playback responses ................... 35 vii List of Figures Figure 1: Sonogram of a Greater Prairie-Chicken call, with several structural elements identified .......................................................................................... 37 Figure 2: Territories of 7 male Greater Prairie-Chickens at the Kreider lek in 2012 near Manhattan, Kansas .......................................................................... 38 Figure 3: Mean differences ± SE in the average boom rate (# booms/min) of male Greater Prairie-Chickens before and during playback treatments .. 39 Figure 4: Mean differences ± SE in the average faceoff rate (# faceoffs/min) of male Greater Prairie-Chickens before and during playback treatments . 40 Figure 5:,,,,,,,,,,,,,l Mean differences ± SE in the proportion of time spent in faceoffs by male Greater Prairie-Chickens before and during playback treatments ...................................................................................... 41 Figure 6: Mean differences ± SE in the closest distance of approach to the playback speaker (m) by male Greater Prairie-Chickens before and during playback treatments ....................................................................... 42 viii Introduction Animal vocalizations play an integral role in signaling aggression, territory defense, and male quality. The acoustic structure of vocalizations is influenced by physical constraints on the vocalizer (Bradbury and Vehrencamp 2011). Thus, physically different individuals may produce different vocalizations. If this is the case, vocalization might be a useful signal for individual recognition as well as allowing for both males and females to assess the fitness of an individual. Auditory signals can also be used to communicate information about spatial location. When an individual’s signals repeatedly originate from a particular location, they may help to define the signaler’s territory (Bee and Gerhardt 2001). Thus, male vocalizations may be a valuable way of establishing territory boundaries. Vocal signaling may also be a mechanism for males to defend their territories by warning other males away before it becomes necessary to escalate to a physically aggressive response. If their territory is encroached upon, males may engage in physical battles with intruders (Apollonio et al. 1992, Arita and Kaneshiro 1985), and those who can successfully maintain their territories may experience higher reproductive fitness (Ryder et al. 2008). Some bird species such
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