DEFINING THE REACTION SPACE OF PREDATOR-PREY INTERACTIONS Ana M. Jurcak A Dissertation Submitted to the Graduate College of Bowling Green State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY May 2018 Committee: Paul Moore, Advisor Emily Freeman Brown Graduate Faculty Representative Jeffrey Miner Shannon Pelini Delbert Smee ii ABSTRACT Paul Moore, Advisor This dissertation contributed to the call for a greater comprehension of sensory ecology within predator-prey interactions, particularly in the non-consumptive effects (NCEs) of predators. I investigated how stimulus modality, predator movement, environmental transmission, prey sensory ecology, pollution, and the interaction of these factors modify prey behavioral responses to predators. Specifically, I experimentally tested three research questions: 1) how the reaction space of predators with different hunting modes in different flow environments altered prey behavior, 2) how modulating signal intensity and prey detection thresholds altered the reaction space, and 3) how the exposure to anthropogenic chemicals altered the reaction space of prey. First, I placed prey (crayfish) in two different environments (flow and no flow) in one of three predator treatments (active predator [bass], sit-and-wait predator [catfish], no predator) and monitored the behavior of the crayfish in a resource patchy environment. Predator hunting mode changed prey behavior, but only in flowing water that would enhance the transmission of predator cues. The most significant interaction between predator treatment and flow environment was found with the active predator in flowing habitats, but this same interaction did not alter NCEs from a sit-and-wait predator. Second, I exposed virile and rusty crayfish to low, medium, or high concentration of odor from largemouth bass and to controls without bass odor and monitored crayfish. The results showed that the behavior of virile crayfish was significantly altered across concentrations more than rusty crayfish, indicating that the virile crayfish may have larger reaction space. Finally, I exposed virile and rusty crayfish to a pesticide (carbaryl) iii then placed the crayfish in a two-choice flume containing predator odor and clean river water to monitor their behavior. I found that the exposure to a carbaryl did not affect the anti-predator behavior of either species. The findings show that each factor of the reaction space is important in understanding and altering NCEs of predators. Additionally, NCEs may be hidden unless the interaction of factors is taken into consideration. Investigating the sensory environment of predator-prey interactions is crucial for better understanding the mechanisms driving the NCEs of predators and their consequences. iv This dissertation is dedicated to my family, John, Maria, and Natalia Jurcak, and Douglas Detter. This would not have been possible without your support through the years. v ACKNOWLEDGMENTS There are many people who I need to thank for their support and guidance through my doctoral degree. First, I want to thank my advisor Dr. Paul A. Moore for the training I’ve received during my six years of being a part of the Lab for Sensory Ecology. Not just my scientific training, but also for all of the advice and guidance on life and beyond. I also want to thank him pushing me to my limits and believing in my abilities. I want to thank my committee members Dr. Emily Freeman, Dr. Jeffrey Miner, Dr. Shannon Pelini, and Dr. Lee Smee, for their guidance, feedback, and advice throughout my degree. I want to thank Dr. Jeffery Miner for agreeing to join my committee in a time of need, Dr. Shannon Pelini for being a positive and strong woman in science role model, Dr. Lee Smee for all of his advice as an expert in the field, and Dr. Emily Freeman Brown for her perspective as someone outside of STEM. I want to thank the past and present members of the Laboratory for Sensory Ecology. Thank you for being a supportive, intellectual, engaging, and hilarious group of people to help me grow as a scientist and person throughout my time in graduate school. My successes during this time would not have happened without all of your support. I would also like to thank Meghan Moore for all of her assistance in set up and data collection for Chapter 3. I would like to thank Bowling Green State University and the Department of Biological Sciences for the infrastructure and funding support throughout my degree. I would like to thank the University of Michigan Biological Station for their funding and infrastructural support over the past five summers of my data collection. All of my research would not have been possible without their generosity. I would also like to thank the Northern Ohio Alliance – Alliance of Graduation Education and the Professoriate for funding and professional development support. Finally, I would like to thank all external funding sources for their contribution to my research. vi Lastly, I would like to thank my family and friends for their love and support throughout my time in the doctoral program. Your constant support has been instrumental in my success. Finally, I would like to thank Douglas Detter for all of his love, support, encouragement, and assistance. You’ve been by my side from day one of this journey and I could not have achieved the success that I have without you by my side. v TABLE OF CONTENTS Page CHAPTER I. GENERAL INTRODUCTION ....................................................................... 1 CHAPTER II. SENSORY SIGNALS AND THE REACTION SPACE IN PREDATOR – PREY INTERACTIONS ....................................................................................................... 7 Introduction ............................................................................................................ 7 Materials and Methods ............................................................................................... 13 Animal collection and holding ....................................................................... 13 Monitoring health and safety of the fish ........................................................ 14 Experimental design ....................................................................................... 14 Experimental arenas ....................................................................................... 15 Shelter and food ............................................................................................. 16 Experimental protocol .................................................................................... 17 Data analysis .................................................................................................. 18 Results ............................................................................................................ 19 Prey behavior in shelter area .......................................................................... 19 Prey behavior in food area ............................................................................. 20 Fish behavior .................................................................................................. 20 Discussion ............................................................................................................ 21 CHAPTER III. DIFFERENT SIZE REACTION SPACES OF TWO PREY SPECIES TO VARIOUS CONCENTRATIONS OF PREDATOR ODOR ............................................... 30 Introduction ............................................................................................................ 30 Materials and Methods ............................................................................................... 33 vi Animal collection and holding ....................................................................... 33 Monitoring health and safety of the fish ........................................................ 34 Experimental design ....................................................................................... 35 Experimental arenas ....................................................................................... 35 Experimental protocol .................................................................................... 36 Food and shelter ............................................................................................. 37 Odor collection and odor prep ....................................................................... 37 Data analysis .................................................................................................. 38 Statistical analysis .......................................................................................... 39 Results ............................................................................................................ 40 Overall effects ................................................................................................ 40 Interaction effects of time spent in each zone ................................................ 40 Interaction effects of time spent stopped in each zone .................................. 41 Interaction effects of walking speed in each zone ......................................... 42 Discussion ............................................................................................................ 43 CHAPTER IV. EFFECT OF ANTHROPGOGENIC CHEMICALS ON THE REACTION SPACE OF PREY ............................................................................................................ 47 Introduction ...........................................................................................................