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Predation and Competition Among Larval Salamanders: The PREDATION AND COMPETITION AMONG LARVAL SALAMANDERS: THE INFLUENCE OF DENSITY DEPENDENCE, PHENOLOGY, FOOD WEB STRUCTURE AND HABITAT COMPLEXITY A Dissertation Presented to the Faculty of the Graduate School at the University of Missouri-Columbia In Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy by THOMAS LEE ANDERSON Dr. Raymond D. Semlitsch, Dissertation Advisor MAY 2016 Dedicated to Raymond D. Semlitsch and Rosemary S. Anderson ACKNOWLEDGEMENTS The number of people that contribute to someone completing a dissertation is innumerable. I owe a great of my development as a scientist to Ray Semlitsch, who pushed me to be better in every possible way. I would not be where I am today without your guidance, advice and support. I will miss the visits to your office to work out some complex problem and instead talking about hunting. Completing this work would not have been possible without the experiences I had prior to graduate school, made possible by Dr. Chris Floyd and the Rocky Mountain Biological Lab, Katie Derr, Dane Cramer and the E.B. Forsythe NWR, and Doug Tempel, Vince Berigan and Sheila Whitmore at Blodgett Forest Station. At Mizzou, this work would not have been possible or nearly as enjoyable without the help of an amazing lab group: Bill Peterman, Grant Connette, Mark Mackey, Britt Ousterhout, Dana Drake, Katie O’Donnell, Jake Burkhart, Freya Rowland, Arianne Messerman, Julia Earl, Rachelle Riegerix, Shannon Pittman, Tom Luhring, Mike Osbourn, and Holly Puglis. All of you made an office space that will be unmatched. I would like thank Ricardo Holdo, Matt Gompper, Debbie Finke and Rex Cocroft for always supporting projects that I was working on, and providing feedback on ideas and manuscripts. John Faaborg, Lori Eggert, and Manuel Leal were great teaching mentors, and the rest of the people on Tucker 2 were great colleagues to be around. Cristina Linares, Katelyn Dodson, Lauren Smith, Josh Robinson, Lauren Hollis, and Chloe Goodson spent many hours helping collect metamorphs, setting up experiments, and doing otherwise mundane task; I am indebted to you all. I thank the many people that contributed to the work at Fort Leonard Wood not presented here but comprised many hours during my dissertation, especially Jennifer Heemeyer, Kim Romine, Kenton ii Lohraff, and the exceedingly pleasant and helpful folks at Range Ops. Many thanks to Howard Whiteman for continued support and a place to crash at Cinnamon, and all the students in his lab at Murray State for reading drafts of manuscripts. The staff and faculty in Tucker, including Nila Emerich, John Walker, Rebecca Ballew, Diane Wyatt, Saadiya Aswad, and Tyeece Little all provided much needed support at various times. I also greatly appreciate Uprise Bakery for allowing me to occupy a table for many hours most Saturday mornings for five years. The Tuesday-Thursday running group helped maintain my sanity over the last five years, especially Katie, Tom, Phil, Nancy, Sam, Laura, Andy, and Ryan. Finally, I thank Mom, Dad and Chris for providing opportunities to go explore the natural world, and for supporting my desires to work in remote areas and study obscure but interesting things. iii TABLE OF CONTENTS Acknowledgements. ....................................................................................................... ii List of Tables.................................................................................................................vii List of Figures ............................................................................................................. viii Abstract .......................................................................................................................... ix Chapter 1. ECOLOGICAL CONTEXT IN PREDATOR-PREY INTERACTIONS Introduction..………………………………………………………………………1 References…………………………………………………………………………4 2. HIGH INTRAGUILD PREDATOR DENSITY INDUCES THINNING EFFECTS ON AND INCREASES TEMPORAL OVERLAP WITH PREY POPULATIONS Abstract……………………………………………………………………………8 Introduction………………………………………………………………………..9 Methods..…………………………………………………………………………12 Results..…………………………………………………………………………..17 iv Discussion..………………………………………………………………………19 Acknowledgements..……………………………………………………………..24 References..………………………………………………………………………25 3. TOP PREDATORS AND HABITAT COMPLEXITY ALTER AN INTRAGUILD PREDATION MODULE IN POND COMMUNITIES Abstract..…………………………………………………………………………37 Introduction..……………………………………………………………………..38 Methods.………………………………………………………………………….41 Results..…………………………………………………………………………..49 Discussion..………………………………………………………………………52 Acknowledgements..……………………………………………………………..58 References..………………………………………………………………………58 4. ASYMMETRIC EFFECTS OF PHENOLOGICAL SHIFTS AND LARVAL DENSITY INFLUENCE INTRAGUILD PREDATION BETWEEN SALAMANDERS Abstract..…………………………………………………………………………76 Introduction..……………………………………………………………………..77 Methods.………………………………………………………………………….80 Results..…………………………………………………………………………..85 v Discussion..………………………………………………………………………89 Acknowledgements..……………………………………………………………..94 References..………………………………………………………………………94 5. SUMMARY Summary..………………………………………………………………………107 APPENDIX A. Non-linear model selection results…………………………………….109 APPENDIX B. Parameter estimates and additional figures for ringed salamanders (Ambystoma annulatum) and spotted salamanders (A. maculatum) ............................... 117 APPENDIX C. Additional tables, figures and statistical analyses of Phenology-Density and Priority Effects experiments .................................................................................. 129 VITA...………………………………………………………………………………….137 vi LIST OF TABLES Chapter 3. Table 1. Test statistics and p-values for ringed salamanders and spotted salamander responses in food web and habitat complexity treatments ................ 68 Chapter 4. Table 1. Posterior distributions of contrasts for ringed salamanders .................. 103 Table 2. Posterior distributions of contrasts for spotted salamanders ................ 104 vii LIST OF FIGURES Chapter 2. Figure 1. Response variables for ringed salamanders .......................................... 27 Figure 2. Response variables for spotted salamanders ........................................ 28 Figure 3. Ringed salamander density and days of temporal overlap with spotted salamanders ........................................................................................... 29 Chapter 3. Figure 1. Diagram of basic IGP module of ringed and spotted salamanders in different top predator and habitat complexity treatments .................................... 55 Figure 2. Response variables of ringed and spotted salamanders among the different top predator and habitat complexity treatments .................................... 56 Figure 3. Response variables of spotted salamanders with ringed salamander survival, top predator and habitat complexity treatments .................................... 57 Chapter 4. Figure 1. Life history diagram and theoretical predictions of phenological shifts in ringed and spotted salamanders ........................................................................ 106 Figure 2. Responses of ringed and spotted salamanders in Phenology-Density experiment ....................................................................................................... 107 Figure 3. Responses of spotted salamanders in Priority Effects experiment ...... 108 viii ABSTRACT The influence of biotic and abiotic factors on species interactions and overall community structure has long interested ecologists. Despite a legacy of interest, there is still ambiguity into the role of biotic and abiotic factors due to highly nuanced, complex networks of interactions that are difficult to comprehend. Yet, understanding how such nuances is an essential goal to determine how they affect population and community structure. Thus, the goal of my dissertation was to understand how multiple biotic and abiotic mechanisms alter interactions among larval stages of two pond-breeding salamanders. Larval stages of pond-breeding salamanders represent an excellent system for understanding how species interactions vary along abiotic and biotic gradients. Intra- and interspecific interactions are frequently determined by size differences among individuals, where larger larvae are predators of smaller larvae and can out-compete them for shared resources. However, when size differences are minimized, only competition occurs. Such conjoined competition and predation is termed intraguild predation, and is a common interaction in many taxa. The factors that determine size differences among individuals (both within and between species) are critical towards to determining both the type of interaction, as well as the strength of such interactions. The focal species I used were the ringed salamander (Ambystoma annulatum) and spotted salamander (A. maculatum). The former breeds earlier than the latter, creating a larval size advantage which results in predation as the dominant interaction between species. However, factors that influence growth rates of ringed salamanders could result in minimized size differences, resulting in a change to the strength or type of interaction that occurs. For my dissertation, I experimentally investigated three different processes that were expected to ix affect the relative importance of predation and competition: density dependence, food web structure, and phenological shifts. In my first chapter, I tested whether the density of ringed salamanders influenced
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