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University of Florida Thesis Or Dissertation Formatting FORAGING ECOLOGY AND CALL RELEVANCE DRIVE RELIANCE ON SOCIAL INFORMATION IN AN AVIAN EAVESDROPPING NETWORK By HARRISON HENRY JONES A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2016 © 2016 Harrison Jones To all those who have given me the inspiration, confidence, and belief to pursue my passion in life ACKNOWLEDGMENTS I thank my parents, who have not only provided unconditional support and encouragement during my degree, but also imbued in me an appreciation for nature and a curiosity about the world. My thanks also go to Elena West, Jill Jankowski, and Rachel Hoang who gave me confidence and the enthusiasm, not to mention sage advice, to apply to a graduate program and follow my passion for ornithology. I would also be remiss to not mention the tremendous support received from the Sieving lab, in particular my amazing officemates Kristen Malone, Willa Chaves, and Andrea Larissa Boesing who were always available to provide help and perspective when needed. I would also like to thank my committee for their helpful input. Katie Sieving was a tremendous help in designing and executing the study, in particular through her knowledge of parid vocalizations. Scott Robinson provided in-depth background knowledge about the study species and insight into the results obtained. And Ben Baiser was invaluable in assisting with the multivariate statistics and generally any other quantitative question I could throw at him. Finally, a big thanks to my many field technicians, Henry Brown, Jason Lackson, Megan Ely, and Florencia Arab, without whom this degree would not be possible. 4 TABLE OF CONTENTS page ACKNOWLEDGMENTS .................................................................................................. 4 LIST OF TABLES ............................................................................................................ 7 LIST OF FIGURES .......................................................................................................... 8 ABSTRACT ..................................................................................................................... 9 CHAPTER 1 INTRODUCTION .................................................................................................... 11 Animal Information Networks .................................................................................. 11 Factors Determining the Value of Social Information to Eavesdroppers ................. 13 2 FORAGING ECOLOGY AND CALL RELEVANCE DRIVE RELIANCE ON SOCIAL INFORMATION IN AN AVIAN EAVESDROPPING NETWORK ............... 17 Methods .................................................................................................................. 17 Study System ................................................................................................... 17 Study Design and Predictions .......................................................................... 19 Characterizing Foraging Behavior .................................................................... 21 Characterizing Sociality and Call Relevance .................................................... 22 Alarm Call Playback Procedures ...................................................................... 23 Data Reduction of Foraging and Microhabitat Variables .................................. 26 Hypothesis Evaluation Using Generalized Linear Models ................................ 27 Results .................................................................................................................... 28 Foraging Observations ..................................................................................... 28 Playback Experiment ........................................................................................ 29 Defining the Microhabitats Occupied During Playback Trials ........................... 31 Defining the Foraging Niches of the Winter Bird Community ............................ 31 Modeling Factors Determining Species’ Reliance on Social Information .......... 33 3 CONCLUDING REMARKS ..................................................................................... 45 The Importance of Foraging Ecology: Aerial Foragers Are Different ...................... 47 Call Relevance Matters, Social System Does Not .................................................. 49 Asymmetric Eavesdropping Networks: Mutualism, Commensalism, or Parasitism? .......................................................................................................... 51 APPENDIX ADDITIONAL TABLES AND FIGURES...................................................... 54 LIST OF REFERENCES ............................................................................................... 65 5 BIOGRAPHICAL SKETCH ............................................................................................ 75 6 LIST OF TABLES Table page 2-1 Predictor variables used in the GLMs for overall (Y/N) response and response type. .................................................................................................... 35 2-2 Summarized foraging data.................................................................................. 36 2-3 Summary of playback response for each species. ............................................. 37 2-4 Model-averaged results of GLMs of overall response and response type. ......... 39 A-1 Principal coordinate axes of the microhabitat measures collected before playback. ............................................................................................................ 54 A-2 Principal coordinate axes from analysis of the foraging ecology data. ............... 54 A-3 Factor loadings for the foraging ecology principal coordinates used in the analysis. ............................................................................................................. 55 A-4 Average values for large and small principal coordinate scores of microhabitat data. ............................................................................................... 56 A-5 Candidate model set of GLMs for overall response used in the model averaging. ........................................................................................................... 56 A-6 Candidate set of best models to explain response type (dive versus freeze). .... 59 A-7 Proportions of foraging maneuver and foraging microhabitat use in a winter bird community. .................................................................................................. 60 7 LIST OF FIGURES Figure page 2-1 Biplots of foraging principal coordinate axes used in the GLM analysis. ............ 41 2-2 Fitted values for the significant predictors of overall response to the titmouse alarm call. ........................................................................................................... 42 2-3 Fitted values for the significant predictors of response type (diving versus freezing response). ............................................................................................. 44 A-1 Biplot of principal coordinate axes Edge-MH and Trunk-MH of the microhabitat variables. ........................................................................................ 64 8 Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science FORAGING ECOLOGY AND CALL RELEVANCE DRIVE RELIANCE ON SOCIAL INFORMATION IN AN AVIAN EAVESDROPPING NETWORK By Harrison Henry Jones August 2016 Chair: Kathryn Sieving Major: Wildlife Ecology and Conservation Vertebrates obtain social information about predation risk by eavesdropping on the alarm calls of sentinel species, which can act as community-wide informants for large numbers of heterospecifics. However, the relative importance of this social information to different eavesdropping species is unknown. We tested the relative importance of four leading hypotheses (foraging ecology, sociality, call relevance based on body size, and local microhabitat) in determining heterospecific reliance on the alarm call of the Tufted Titmouse (Baeolophus bicolor) in a Florida winter bird community. We presented 16 forest species, exhibiting broad social, ecological, and taxonomic variability, with a titmouse alarm call (known to generate strong anti-predator responses in birds) and quantified responses to playback. Predictor metrics representing local microhabitat were assessed during the playback study, whereas species’ foraging ecology and sociality were determined through independent behavioral observations in the study area. Following data reduction procedures applied to 56 total measures, we tested for the effects of 11 final predictor variables (expressing four hypothesized factors plus nuisance variables) on two different response metrics (presence and type of escape response). Using generalized linear modeling we determined best predictors 9 using model averaging of a candidate set of models (ΔAICc < 2). Overall response was best predicted by foraging ecology and call relevance; relatively larger-bodied canopy foragers and those that use more aerial maneuvers responded less often. Escape behavior type was best explained by a species’ foraging distance from the tree trunk; near-trunk foragers were more likely to freeze in response to playback, whereas foliage gleaners spending time far from trunks were more likely to dive for cover. Our work clearly
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