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Ecology of the Eastern Long-Tailed Salamander (Eurycea

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Ecology of the Eastern Long-Tailed Salamander (Eurycea ECOLOGY OF THE EASTERN LONG-TAILED SALAMANDER (EURYCEA LONGICAUDA LONGICAUDA) ASSOCIATED WITH SPRINGHOUSES by Nathan H. Nazdrowicz A dissertation submitted to the Faculty of the University of Delaware in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Entomology and Wildlife Ecology Spring 2015 © 2015 Nathan H. Nazdrowicz All Rights Reserved ProQuest Number: 3718365 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. ProQuest 3718365 Published by ProQuest LLC (2015). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code Microform Edition © ProQuest LLC. ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, MI 48106 - 1346 ECOLOGY OF THE EASTERN LONG-TAILED SALAMANDER (EURYCEA LONGICAUDA LONGICAUDA) ASSOCIATED WITH SPRINGHOUSES by Nathan H. Nazdrowicz Approved: __________________________________________________________ Jacob L. Bowman, Ph.D. Chair of the Department of Entomology and Wildlife Ecology Approved: __________________________________________________________ Mark W. Rieger, Ph.D. Dean of the College of Agriculture and Natural Resources Approved: __________________________________________________________ James G. Richards, Ph.D. Vice Provost for Graduate and Professional Education I certify that I have read this dissertation and that in my opinion it meets the academic and professional standard required by the University as a dissertation for the degree of Doctor of Philosophy. Signed: __________________________________________________________ Jacob L. Bowman, Ph.D. Professor in charge of dissertation I certify that I have read this dissertation and that in my opinion it meets the academic and professional standard required by the University as a dissertation for the degree of Doctor of Philosophy. Signed: __________________________________________________________ Douglas W. Tallamy, Ph.D. Member of dissertation committee I certify that I have read this dissertation and that in my opinion it meets the academic and professional standard required by the University as a dissertation for the degree of Doctor of Philosophy. Signed: __________________________________________________________ Christopher K. Williams, Ph.D. Member of dissertation committee I certify that I have read this dissertation and that in my opinion it meets the academic and professional standard required by the University as a dissertation for the degree of Doctor of Philosophy. Signed: __________________________________________________________ Don C. Forester, Ph.D. Member of dissertation committee I certify that I have read this dissertation and that in my opinion it meets the academic and professional standard required by the University as a dissertation for the degree of Doctor of Philosophy. Signed: __________________________________________________________ Mac F. Given, Ph.D. Member of dissertation committee ACKNOWLEDGMENTS I express my sincere appreciation to my advisor, Dr. Jacob L. Bowman, and my other committee members, Dr. Christopher K. Williams, Dr. Doug W. Tallamy, Dr. Don C. Forester, and Dr. Mac F. Given for their knowledge and guidance on this project. I express my sincere gratitude to all the property owners who granted me access to explore their springhouses. In particular, I thank Mike and Angie Riska, Fred and Cindy Von Czoernig, Bob Deen, Alan Crawford, Dale Longmaid, Brandywine Conservancy, Tri- State Bird Rescue, White Clay Creek State Park, and Pennsylvania Resource Council for allowing me access to their springhouses at night through early morning hours every few weeks for several years. I also thank Kevin Fryberger, who assisted with searching for springhouse and property access in Pennsylvania, and Sam and Helen Cupp, who shared their knowledge on springhouse in Delaware and provided a copy of extensive notes on springhouses. I am grateful to all those who offered advice or assisted with data collection in the field, in particular P. Adkins, J. Alexander, L. Allison, B. Bahder, C. Bartlett, E. Brazell, R. Bowker, B. Brown, M. Cacciapaglia, G. Colligan, S. T. Dash, L. Deaner, A. DeWire, M. DiBona C. Diggins, R. Donovall, D. Egan, T. Eisenhower, J. Evans, E. Farris, R. Fausnaught, R. Franz, A. Griger, E. Groth, A. Gonzon, S. Haldeman, B. Harvath, A. Hill, K. Howard, C. Howey, A. Hulse, R. Iglay, B. Jennings, M. Johnston, M. Jonas, S. Kuebbing, E. Lake, V. Maruschak, M. Miller, R. Misiewicz, D. Nonne, A. Pisano, R. Plank, N. Ranalli, C. Rhoads, J. Rogerson, R. Sawyer, M. Scafini, F. Schmid, A. Schneider, K. Stocki, D. Stoner, J. Thompson, K. Wallace, K. Ward, S. Williamson, G. Wirth, R. Wall, N. Watson, J. Yeager, and M. Zuefle, I also thank Jim White for v assistance with data collection, driving to field sites, and providing input on sample design. Finally, I would like to thank Richard Franz for discussing his observations of eastern long-tailed salamanders in Maryland. vi TABLE OF CONTENTS LIST OF TABLES ......................................................................................................... ix LIST OF FIGURES ..................................................................................................... xii ABSTRACT .................................................................................................................. xv 1 INTRODUCTION .............................................................................................. 1 Study Areas ......................................................................................................... 5 2 LIFE HISTORY AND ECOLOGY OF EASTERN LONG-TAILED SALAMANDERS (EURYCEA LONGICAUDA LONGICAUDA) ASSOCIATED WITH SPRINGHOUSES ....................................................... 19 Introduction ....................................................................................................... 19 Methods............................................................................................................. 21 Site Visits .................................................................................................... 21 Reproductive Data ...................................................................................... 22 Mark-Recapture Data .................................................................................. 24 Mark-Recapture Analysis ........................................................................... 28 Results ............................................................................................................... 31 Reproduction ............................................................................................... 31 Courtship and Mating ........................................................................... 31 Reproduction ......................................................................................... 32 Egg Development.................................................................................. 36 Larval Period ............................................................................................... 37 Activity Patterns.......................................................................................... 39 Daily Activity........................................................................................ 39 Seasonal Activity .................................................................................. 40 Mark-Recapture .......................................................................................... 42 Captures ................................................................................................ 43 Age Composition .................................................................................. 45 Sex Composition ................................................................................... 45 vii Recaptures ............................................................................................. 46 Population Estimates ............................................................................. 47 Discussion ......................................................................................................... 48 Reproduction ............................................................................................... 48 Larval Period ............................................................................................... 57 Activity Patterns and Movements ............................................................... 59 Reproduction ......................................................................................... 60 Overwintering ....................................................................................... 63 Foraging ................................................................................................ 64 Population Dynamics .................................................................................. 66 Age and Sex Composition .................................................................... 68 Sex Ratios ............................................................................................. 69 Population Trends ................................................................................. 72 3 FACTORS INFLUENCING SITE OCCUPANCY OF
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