The Population Ecology and Behavior of the Cave Salamander, Eurycea Lucifuga (Rafinesque, 1822)
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University of Louisville ThinkIR: The University of Louisville's Institutional Repository Electronic Theses and Dissertations 8-2018 The population ecology and behavior of the cave salamander, Eurycea lucifuga (Rafinesque, 1822). Joseph Gavin Bradley University of Louisville Follow this and additional works at: https://ir.library.louisville.edu/etd Part of the Behavior and Ethology Commons, Population Biology Commons, and the Zoology Commons Recommended Citation Bradley, Joseph Gavin, "The population ecology and behavior of the cave salamander, Eurycea lucifuga (Rafinesque, 1822)." (2018). Electronic Theses and Dissertations. Paper 3041. https://doi.org/10.18297/etd/3041 This Doctoral Dissertation is brought to you for free and open access by ThinkIR: The University of Louisville's Institutional Repository. It has been accepted for inclusion in Electronic Theses and Dissertations by an authorized administrator of ThinkIR: The University of Louisville's Institutional Repository. This title appears here courtesy of the author, who has retained all other copyrights. For more information, please contact [email protected]. THE POPULATION ECOLOGY AND BEHAVIOR OF THE CAVE SALAMANDER, EURYCEA LUCIFUGA (RAFINESQUE, 1822) By Joseph Gavin Bradley B.S., University of Louisville, 2011 M.S., University of Louisville, 2016 A Dissertation Submitted to the Faculty of the College of Arts and Sciences of the University of Louisville In Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in Biology Department of Biology University of Louisville Louisville, Kentucky August 2018 THE POPULATION ECOLOGY AND BEHAVIOR OF THE CAVE SALAMANDER, EURYCEA LUCIFUGA (RAFINESQUE, 1822) By J. Gavin Bradley B.S., University of Louisville, 2011 M.S., University of Louisville, 2016 A Dissertation Approved on July 3, 2018 by the following Dissertation Committee: Dissertation Director Perri Eason James Alexander Julian Lewis William Pearson Steve Yanoviak ii DEDICATION To Emma, Jameson, and Bailey iii ACKNOWLEDGEMENTS I would like to thank, tremendously, Dr. Perri Eason for her expert guidance and having the patience for my independence (and perhaps stubbornness at times) throughout this entire process. To Dr. Julian Lewis, I give thanks to his great devotion of knowledge and time in the field at the many caves we visited over these few years. I am further grateful for the opportunities, second chances, and skillful instruction of Dr. Steve Yanoviak, Dr. William Pearson, and Dr. James Alexander. Much of this research would not have been possible without the correspondence and help from Clay Foreman and the staff at E.P. “Tom” Sawyer State Park. Thanks to Svend Bentsen for producing historical documents of Sauerkraut Cave. I thank Allen Pursell and Mike Everidge of The Nature Conservancy for their time in showing and granting me access to study sites in Indiana. I thank Gary Roberson and the staff at Indiana Caverns for their friendliness, accommodation, and willingness to allow me to explore research opportunities. I would like to recognize Ken Bailey for his time and efforts in the field as well, and for introducing me to many Kentucky caves. I had an amazing group of undergraduates that assisted me with data collection. I am indebted to Paige Wilson and Danica Shepherd for their efforts and long-term dedication to this project. I much appreciate Faith Bowers for taking on the reigns after Paige and Danica moved on. I thank Logan Stone, Nick Callahan, and Will Seibt for their extended efforts. Emily Petri and Isaac Kaufman came on in the closing months of this iv project, and I am grateful for their help in the field and the lab. For any of those I have failed to mention, know you are appreciated. Many fellow graduate students helped me in various ways during this process. I thank Dr. Benjamin Adams, Dr. Evan Gora, and Lindsay Nason for statistical guidance. To past and present Eason lab members (Dr.’s Carl Cloyed, Sarah Fauque, Micah Perkins, Bill Persons, Victoria Prescott, Piyumika Suriyampola, and Jared Wood; Lindsay Nason and Jeeva Rathnaweera) and Megan DeWhatley, thank you all for your time, comments, suggestions, and help in the field. To Emma Bradley, never enough can be said. Thank you for your love, willingness, patience, support, and motivation toward my achievements. To the rest of my family, thank you for the love and support. This research was approved by the University of Louisville’s Institutional Animal Care and Use Committee (IACUC #17005), Kentucky Department of Fish and Wildlife Resources (permit #SC1511236, #SC1611045, and #SC1711045), Kentucky Department of Parks (permit #1412 and #1701), and the Indiana Department of Natural Resources: Division of Nature Preserves (permit #NP17-52). v ABSTRACT THE POPULATION ECOLOGY AND BEHAVIOR OF THE CAVE SALAMANDER, EURYCEA LUCIFUGA (RAFINESQUE, 1822) J. Gavin Bradley July 3, 2018 The Cave Salamander, Eurycea lucifuga (Rafinesque, 1822), is a little-known species, yet a common inhabitant of caves in the eastern United States. Salamanders are often important components of ecological communities and ecosystems, influencing critical processes such as nutrient cycling and community composition through their predation on invertebrates. Cave-dwelling salamanders such as E. lucifuga may thus appreciably influence the relatively simple ecosystems and communities of caves. Any such influence may be particularly important because these habitats and the organisms that reside in them are often of conservation concern. I used non-invasive methods to study the demographics, movements, and habitat selection of E. lucifuga at Sauerkraut Cave in Louisville, Kentucky. I also conducted an experimental manipulation using clay models to test predation risk to Cave Salamanders in caves and forests in southern Indiana. I discovered that E. lucifuga have consistent and distinguishable spot patterns that can be used to identify individuals. Populations of this species may be much larger than previously thought, potentially contributing relatively large and seasonally variable biomass to spring cave systems. Furthermore, this species migrates seasonally within vi caves using abiotic environmental cues that indicate seasonal change, potentially shuttling acquired energy from forests to deep underground. I also demonstrated that Cave Salamanders likely use caves, and particularly cave walls, as a refuge from greater potential predation risk in forests. This research provides much-needed information on this species and is suggestive that cave-dwelling salamanders may have important ecological roles in subterranean environments. vii TABLE OF CONTENTS PAGE ACKNOWLEDGEMENTS ....................................................................................................... iv ABSTRACT ......................................................................................................................... vi LIST OF TABLES .................................................................................................................. ix LIST OF FIGURES ..................................................................................................................x CHAPTER I: DISSERTATION INTRODUCTION .........................................................................1 CHAPTER II: IMAGE-BASED METHODS FOR THE IDENTIFICATION AND MEASUREMENT OF CAVE SALAMANDERS, EURYCEA LUCIFUGA (RAFINESQUE, 1822) ............................... 11 INTRODUCTION ...................................................................................................... 11 METHODS .............................................................................................................. 12 RESULTS ................................................................................................................ 22 DISCUSSION ........................................................................................................... 29 CHAPTER III: OBSERVATIONS ON THE POPULATION ECOLOGY OF THE CAVE SALAMANDER, EURYCEA LUCIFUGA (RAFINESQUE, 1822) ............................................ 35 INTRODUCTION ...................................................................................................... 35 METHODS .............................................................................................................. 37 RESULTS ................................................................................................................ 41 DISCUSSION ........................................................................................................... 46 CHAPTER IV: THE EFFECTS OF CLIMATE AND BEHAVIOR ON MIGRATORY MOVEMENTS OF CAVE SALAMANDERS, EURYCEA LUCIFUGA (RAFINESQUE, 1822) ................................ 55 INTRODUCTION ...................................................................................................... 55 METHODS .............................................................................................................. 60 RESULTS ................................................................................................................ 66 DISCUSSION ........................................................................................................... 77 CHAPTER IV APPENDIX .......................................................................................... 86 CHAPTER V: REFUGE FROM DANGER: HABITAT SELECTION BY CAVE SALAMANDERS, EURYCEA LUCIFUGA (RAFINESQUE, 1822) .................................................................... 88 INTRODUCTION .....................................................................................................