MORPHOLOGY, ECOPHYSIOLOGY, AND IMPACTS OF NONINDIGENOUS POMACEA IN FLORIDA By JENNIFER L. BERNATIS A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2014 1 UMI Number: 3647807 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. UMI 3647807 Published by ProQuest LLC (2014). Copyright in the Dissertation held by the Author. Microform Edition © ProQuest LLC. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, MI 48106 - 1346 © 2014 Jennifer L. Bernatis 2 This is for all of the grad students who think the process will never end. And to Tabitha and Snuffy, you are truly missed but will never be forgotten. 3 ACKNOWLEDGMENTS I would like to thank my committee, Dr. Steve Johnson (Chair), Dr. Mark Brenner (Co-chair), Dr. Ken Langeland, Dr. Jim Williams, and Dr. Tom Frazer for their support and patience through this process. I am also very thankful to Dr. Chad Cross, Environmental and Occupational Health, University of Nevada Las Vegas, and Dr. Erin Leone, Florida Fish and Wildlife Conservation Commission, for invaluable statistical help and support. In addition, I would like to thank Mr. Gary Warren, Florida Fish and Wildlife Conservation Commission, and Dr. Iain McGaw, Memorial University, for their continued support and encouragement throughout this project. I would also like to thank Mr. Bill Coleman, Florida Fish and Wildlife Conservation Commission, for providing the resources for much of this project. And finally, I would like to thank my parents, Robert and Terry Bernatis, for not minding the fact they have a professional student for a daughter. 4 TABLE OF CONTENTS page ACKNOWLEDGMENTS .................................................................................................. 4 LIST OF TABLES ............................................................................................................ 8 LIST OF FIGURES .......................................................................................................... 9 ABSTRACT ................................................................................................................... 10 CHAPTER 1 INTRODUCTION .................................................................................................... 12 Background ............................................................................................................. 12 Morphological Characteristics ................................................................................. 13 Biology .................................................................................................................... 15 Habitat and Environmental Tolerances ............................................................ 15 Feeding ............................................................................................................ 16 Reproduction .................................................................................................... 17 Age and Growth ............................................................................................... 19 Pathways of Introduction ......................................................................................... 20 Effects of Introduction ............................................................................................. 21 Summary and Research Goals ............................................................................... 24 2 MORPHOLOGY ...................................................................................................... 32 Background ............................................................................................................. 32 Methods .................................................................................................................. 35 Collection and Holding Procedures .................................................................. 35 Measuring Procedures ..................................................................................... 36 Statistical Analysis ............................................................................................ 37 Results .................................................................................................................... 38 Nonparametric MANOVA ................................................................................. 38 Preliminary Canonical Discriminant (CDF) and Discriminant Function Analyses........................................................................................................ 40 Forward Stepwise Analysis .............................................................................. 43 Follow-up Canonical Discriminant and Discriminant Function Analyses .......... 44 Pre and Post Cluster Frequencies .................................................................... 46 Discussion .............................................................................................................. 47 3 PHYSIOLOGICAL TOLERANCES ......................................................................... 67 Background ............................................................................................................. 67 Methods .................................................................................................................. 71 Collection and Holding ..................................................................................... 71 5 General Procedures ......................................................................................... 72 Starvation ......................................................................................................... 73 Salinity .............................................................................................................. 73 pH ..................................................................................................................... 74 Desiccation ....................................................................................................... 74 Statistical Analyses .......................................................................................... 76 Results .................................................................................................................... 76 Starvation ......................................................................................................... 76 Salinity .............................................................................................................. 77 pH ..................................................................................................................... 78 Desiccation ....................................................................................................... 79 Discussion .............................................................................................................. 81 4 FEEDING RATES AND PREFERENCES ............................................................... 93 Background ............................................................................................................. 93 Methods .................................................................................................................. 96 General Procedures ......................................................................................... 96 Feeding Rates .................................................................................................. 97 Feeding Preferences ........................................................................................ 98 Single age group ........................................................................................ 98 Mixed age group ........................................................................................ 99 Statistical Analysis ............................................................................................ 99 Results .................................................................................................................. 100 Feeding Rates ................................................................................................ 100 Feeding rates of adult P. canaliculata ...................................................... 100 Feeding rates of adult P. maculata .......................................................... 101 Feeding rates of juvenile P. maculata ...................................................... 102 Comparison of feeding rates of adult snails ............................................. 103 Comparisons of feeding rates of P. maculata age groups ....................... 103 Feeding Preferences ...................................................................................... 104 Six-week feeding preference study .......................................................... 104 Two-week feeding preference trials ......................................................... 106 Discussion ............................................................................................................ 107 5 EFFECTIVENESS OF MANUAL REMOVAL FOR CONTROL OF NONINDIGENOUS APPLE