Vocal Behavior Is Correlated with Mixed-Species’ Reactions to Novel Objects
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University of Tennessee, Knoxville Trace: Tennessee Research and Creative Exchange Doctoral Dissertations Graduate School 5-2015 Mixed-species Flock Members’ Reactions to Novel and Predator Stimuli Sheri Ann Browning University of Tennessee - Knoxville, [email protected] Recommended Citation Browning, Sheri Ann, "Mixed-species Flock Members’ Reactions to Novel and Predator Stimuli. " PhD diss., University of Tennessee, 2015. https://trace.tennessee.edu/utk_graddiss/3327 This Dissertation is brought to you for free and open access by the Graduate School at Trace: Tennessee Research and Creative Exchange. It has been accepted for inclusion in Doctoral Dissertations by an authorized administrator of Trace: Tennessee Research and Creative Exchange. For more information, please contact [email protected]. To the Graduate Council: I am submitting herewith a dissertation written by Sheri Ann Browning entitled "Mixed-species Flock Members’ Reactions to Novel and Predator Stimuli." I have examined the final electronic copy of this dissertation for form and content and recommend that it be accepted in partial fulfillment of the requirements for the degree of Doctor of Philosophy, with a major in Psychology. Todd M. Freeberg, Major Professor We have read this dissertation and recommend its acceptance: Gordon M. Burghardt, Matthew Cooper, David A. Buehler, Kathryn E. Sieving Accepted for the Council: Dixie L. Thompson Vice Provost and Dean of the Graduate School (Original signatures are on file with official student records.) Mixed-species Flock Members’ Reactions to Novel and Predator Stimuli A Dissertation Presented for the Doctor of Philosophy Degree The University of Tennessee, Knoxville Sheri Ann Browning May 2015 i Copyright © 2015 by Sheri Ann Browning All rights reserved. ii ACKNOWLEDGEMENTS Thank you to all the bird lovers in my family, including Mom, Dad, Nana, and Grandmother, who taught me at a young age to appreciate birds and their predators. Who could have known that all the time I spent watching the backyard birds and the occasional hawk were preparing me for this 8-year journey? Mom and Dad – I love you both very much, and would not have made it to where I am now without your love and support. Thank you to my brother, Bobby, and sister, Holly. Both have supported me, even without truly understanding what I was studying or why. That’s what I call love! Of course, many thanks go to my advisor, Dr. Todd Freeberg. I have felt like Sisyphus at times, struggling to push this dissertation boulder up the (Smoky) Mountains, but he has always been there to help, (de)motivate, and provide advice. I can only hope to be as accomplished as he is in the future. Thanks also to my committee members, Drs. Gordon Burghardt, Matthew Cooper, David Buehler, and Katie Sieving, who have supported me, sparked interesting conversations, and provided constructive criticism throughout my journey. Because of them, I have grown into a better thinker, researcher, and writer. I am also thankful for my lab mates, David Book, Steven Kyle, and Brittany Coppinger, and NIMBioS colleagues, Arik Kershenbaum, Amiyaal Ilany, and Elizabeth Hobson, who have always been there when I needed help, whether it be catching rogue birds in the aviaries, taking care of the birds, or providing feedback on manuscripts or presentations. Thanks also go to Suzanne Winters and Carrie Newton-Hodge for their help with data collection and inter-rater reliability during the initial experiment that sparked my interest in mixed-species flock members’ reactions to novel stimuli. I am also grateful to Megan Wu and Phylicia Washington for their help with inter-rater reliability for subsequent experiments. iii I am appreciative of Rosalee Kashel and Bailey Mulderig, who took care of my birds for me this past Christmas, allowing me a short, but glorious break from fieldwork. Special thanks to Kevin Hoyt and the staff at the University of Tennessee Forest Resources AgResearch and Education Center. I spent so much time there, that I sometimes felt like it would have been more efficient if I just set up camp next to the aviaries for the winter! Thanks to the University of Tennessee’s Psychology Department for their dissertation grant and also to Connie Ogle, who has been able to help me when I needed it, even when she was being pulled in multiple directions. Thanks to all of my friends in Tennessee and Virginia who have cheered me on as I trudged through seemingly endless files to code, wished me warm thoughts on especially cold data collection days, provided fun incentives if I was productive, and allowed me to complain to them when I had birdless trapping days. And last, but not least, thanks go to all the chickadees, titmice, and nuthatches I have observed, trapped, banded, and/or studied. I couldn’t have done it without them! They are smarter than I give them credit for (and I give them lots of credit!) They are also stubborn and tested my patience more often than I would have liked, but it was worth it to get a better understanding of what goes on in their world. Dr. Seuss’s character, the Lorax, says he speaks for the trees. In my dissertation, I’m speaking for the birds. iv ABSTRACT Novel stimuli are ubiquitous. Few studies have examined mixed-species group reactions to novelty, although the complex social relationships that exist can affect species’ behavior. Additionally, studies rarely consider possible changes in communication. However, for social species, changes in communication, including rates, latencies, or note-types within a call, could potentially be correlated with behavioral traits. As such, this research aimed to address whether vocal behavior is correlated with mixed-species’ reactions to novel objects. I first tested the effect of various novel stimuli on the foraging and calling behavior of Carolina chickadees, Poecile carolinensis, and tufted titmice, Baeolophus bicolor. Chickadees and titmice both had longer latencies to forage in the presence of novel stimuli. Chickadees also modified their vocal behavior, having shorter latencies to call and using more ‘D’ notes in their calls in the presence of novel stimuli compared to titmice. Chickadees and titmice reacted to the novel stimuli similarly to how I would expect them to react to a predator. Therefore, a second experiment was conducted directly comparing chickadee and titmouse reactions to a novel (Mega Bloks®) stimulus and a predator (Cooper’s hawk) stimulus. Chickadees and titmice had an intermediate latency to forage in the presence of a novel stimulus compared to control and predator contexts. Again, chickadees had shorter calling latencies across contexts compared to titmice. As a final experiment, using semi-naturalistic aviaries, I tested whether chickadee flock size and the presence or absence of titmice influenced reactions to novel and predator stimuli. Chickadees called more in smaller chickadee flocks compared to larger chickadee flocks, and also when titmice were absent compared to when they were present. These results were stronger in predator contexts compared to novel contexts. This suggests that conspecific flock size influences calling behavior, such that smaller flocks, which may experience higher stress levels and may be required to exhibit more anti-predatory behavior, call more than larger flocks. Taken together, this work has important v implications for the complexity of social relationships in mixed-species groups, the social roles species play within the group, and how group size influences vocal behavior and reactions to various degrees of threat. vi TABLE OF CONTENTS CHAPTER I: INTRODUCTION ................................................................................... 1 Scope ...................................................................................................................... 1 Novel Stimuli, Neophobia, and Neophilia ..........................................................2 Personality Terminology ...................................................................................... 5 History of Personality and Novelty Research in Animals ..................................9 How to Measure Neophobia and Neophilia ...................................................... 12 Habituation and Dishabituation ........................................................................ 14 Influences on Neophobia and Neophilia ........................................................... 15 Vocal Behavior: Another Possible Novelty Response Measure...................... 22 The Study System ............................................................................................... 24 Outline of this Dissertation Work..................................................................... 26 Appendix: Tables ................................................................................................... 28 CHAPTER II The differential reactions of Carolina chickadees, Poecile carolinensis, and tufted titmice, Baeolophus bicolor, to novel stimuli ............. 31 Abstract................................................................................................................... 32 Introduction ........................................................................................................... 33 Predictions ...........................................................................................................35 Stimuli.................................................................................................................. 37 Methods .................................................................................................................