ELIMIA CURVICOSTATA SPECIES-COMPLEX WITHIN THE RIVER DRAINAGES OF THE SOUTHEASTERN UNITED STATES: MORPHOLOGY, DNA, AND BIOGEOGRAPHY BY ELIZABETH LOUISE MIHALCIK 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 1998 Copyright 1998 by Elizabeth Louise Mihalcik This is dedicated to my dogs Annie, Buffy, Darwin, Benjamin, Nickie, and Gina who accompanied me on this journey. ACKNOWLEDGMENTS chairman, for I would like to thank Dr. F. Wayne King, my committee sponsoring me as a doctoral student in the Department of Wildlife Ecology and Conservation. F. Dr. I would like to thank the members of my committee, Dr. John Eisenberg, Patricia Werner, Dr. Brian Bowen, and Dr. David Dilcher. I would like to thank Dr. Fred G. Thompson for all his enthusiasm, advice, encouragement, and assistance with this project. I would like to thank Ms. Anna Clark, Ms. Anna Bass, Ms. Alicia Francisco, Dr. William Farmerie, and Ms. Angela Garcia-Roderguiez of Molecular Services for their time, patience, expertise, and knowledge. I would like to thank Dr. David Moraga, ICBR Educational Core, for allowing me to use the laboratory and to store my specimens in the -80°C Ultrafreezer. I would like to thank Dr. Frank Nordlie for the use of his laboratory and equipment to perform the limnological analyses for this investigation. I would like to thank Dr. Steve Karl, University of South Florida, and Dr. Charles Lydeard, University of Alabama, for their assistance on the DNA analyses. I would like to thank Dr. John B. Burch, Curator of the Mollusk Division at the University of Michigan Museum of Zoology, for supporting me through my program, loaning the type material, and allowing me to visit the collection. iv Hershler, Curator of Mollusks at the National I would like to thank Dr. Robert Museum of Natural History, for loaning the types designated by Lea. Curator of Mollusks at the British Museum I would like to thank Mr. Fred Naggs, of Natural History, for loaning the types designated by Reeve. Curator of the Mollusk Division, at I would like to thank Dr. Kenneth Boss, Harvard University, for allowing me to visit the collection. Biologist at the Florida Caverns State I would like to thank Mr. Mark Ludlow, Park, for assisting me with the collecting permit for the park. Department of Game and Fish I would like to thank Dr. Paul Moler, Florida Commission, for collecting snails in the Florida Panhandle for this study. Florida Museum of Natural I would like to thank the Computer Staff of the History, Mr. Dick Ruble, Mr. Sean Thompson, and Mr. Bill Payne, for all their expertise and assistance. for his assistance with I would like to thank Mr. Jay Harrison, IFAS Statistics, writing programs for SAS and the data analyses of this project. visit to I would like to thank Ms. Renee Sherman for assistance with my the Mollusk Division at the University of Michigan. I would like to thank Mr. Daniel Graf for sharing his knowledge on the history of malacology relating to pleurocerids and his assistance with my visit to the Mollusk Division at the Museum of Comparative Zoology at the Harvard University. I would like to thank Mr. Louis Clark, Mr. Roger Hoover, Ms. Dale Johnson, and Mr. Jeff Knee for the illustrations and figures to accompany this document. I would like to thank the American Museum of Natural History for awarding me the Theodore Roosevelt Memorial Fund which allowed me to travel to the rivers in Georgia. v Comparative Zoology for awarding me the I would like to thank the Museum of research collection. Ernst Mayr Award so I could visit the mollusk Protection Agency for awarding me a I would like to thank the US Environmental investigation. STAR Graduate Research Fellowship so I could finish this (sister), Lori M. Bishop (niece), I would like to thank my family, Joan Bishop Randy W. Bishop (nephew) for all their help and support. Ms. Joanne M. I would like to thank my close friends, Ms. Stevie A. Rutter, Yonge, Ms. Charlene Fronziak, Ms. Mary Lou Lyman, Mr. Jack A. Rusher, Ms. Katherine L. Roth, and Ms. Joan Fremo for their emotional support. for her many words of I owe a great deal to Ms. Joanne C. Yonge (1945-1996) encouragement when I needed them the most to pursue this program. vi TABLE OF CONTENTS ACKNOWLEDGMENTS IV LIST OF TABLES x LIST OF FIGURES xii ABSTRACT xiv CHAPTER 1 INTRODUCTION 1 Taxonomic Overview 1 Historical Background 2 The Problems associated with Taxonomic Studies 11 The Taxonomic Focus of this Study 13 Study Objectives and Implications 14 2 MATERIALS AND METHODS 16 Field Collection Methodologies 16 Animal Description 16 The Study Area 17 Field Collection 18 Data Collection 19 Shell Morphological Analyses 19 Examination of Relevant Museum Types 19 Shell Morphometries 20 Genetic Analysis 23 Mitochondrial DNA Isolation 23 Polymerase Chain Reaction (PCR) Methodology 25 Agarose Gel Purification 25 DNA Sequencing 26 Limnology 26 Water Collection 26 Water Quality Analyses 27 Statistical Analyses 28 Statistical Analysis of Shell Characters 28 Statistical Analyses of the Mitochondrial DNA 28 vii 3 LABORATORY AND FIELD RESULTS 46 Interspecific Variation of the Southeastern Elimia 46 Shell Morphological Analysis 46 Cluster Analysis using Shell Morphology 54 Genetic Analysis of the Cytochrome oxidase subunit 1 57 Sequence Divergence 57 Relationships of C01 Genotypes 65 Intraspecific Variation of the Southeastern Elimia 70 Shell Morphological Analysis 70 Analysis of Types 56 Species and their Geographic Distribution 80 Limnological Analyses 81 4 SYSTEMATICS 86 Overview of the Gulf Coastal fauna 87 Synopses of the Gulf Coastal Rivers Species 89 Overview of Altamaha River fauna 102 Synopses of the Altamaha River Species 102 5 DISCUSSION 109 Regional Physiography 109 The Physiography of Georgia Rivers 111 The Physiography of Alabama Rivers 111 The Physiography of Florida Rivers 113 Unraveling Taxonomic Problems 1 14 Shell Morphology vs mitochondrial DNA 114 Biodiversity and Biogeography of the Southeastern Rivers 115 Elimia Biodiversity and Faunal Distribution 115 Biodiversity of other Southeastern Freshwater Groups 124 Biogeography of the other Southeastern Groups 128 Origin, Dispersal, and Evolution of the Region's Fauna 130 Drainage Evolution within the Region 133 Evolution of the Southeastern Freshwater Fauna 135 Biodiversity Loss and Conservation Implications 137 The Decline of Aquatic Mollusks 137 Threats to the Aquatic Ecosystems 138 Biodiversity Conservation 140 Habitat Conservation 33 APPENDICES A VOUCHER SPECIMEN INFORMATION 145 B MORPHOMETRIC ANALYSES OF THE SOUTHEASTERN ELIMIA USED IN THE CANONICAL DISCRIMINANT ANALYSIS 155 viii C MORPHOMETRIC ANALYSES OF THE TYPES RELEVANT TO THE SOUTHEASTERN ELIMIA 178 D LIMNOLOGICAL DATA OF THE FIELD LOCALITIES OF THE SOUTHEASTERN ELIMIA 189 REFERENCES I 93 BIOGRAPHICAL SKETCH 209 ix LIST OF TABLES Table page 1 . Original Taxonomic Designations of southeastern Elimia 5 2. Species recognized by Tryon, Jr 7 3. Species recognized by Goodrich 8 4. Species recognized by Clench and Turner 9 5. Species recognized by Burch 10 6. Species recognized by Chambers 12 7. Localities sampled within the lower Alabama River drainage system 31 8. Localities sampled within the Escambia River drainage systems 33 9. Localities within the Choctawhatchee and Yellow River drainage systems 37 10. Localities sampled within the Apalachicola River drainage system 40 1 1 . Localities sampled within the Atlantic Coastal river drainage systems 44 12. Summary of Analyses performed within the southeastern river systems 47 13. ANOVAS of the 12 Shell Characters 49 14. Results of the Canonical Discriminant Analysis 49 15. Standardized Canonical Coefficients 49 16. Pairwise Distance Matrix for Cluster Analysis 55 17. Nucleotide Sequence Divergence within the southeastern Elimia 71 18. Morphological Characteristics of the southeastern Elimia 74 19. Summary of limnological analyses associated with Elimia species 85 20. Review of Taxonomic names associated with Elimia densicostata 91 x Distribution of selected freshwater taxa within the southeastern river systems xi LIST OF FIGURES Figure page 1 . North American Elimla species 3 2. Elimia species 21 3. Localities sampled within the Lower Alabama River drainage system 30 4. Localities sampled within the Escambia River drainage system 32 5. Localities sampled within the Choctawhatchee River drainage system 36 6. Localities sampled within the Apalachicola River drainage system 39 7. Localities sampled within the Atlantic Coastal river drainage systems 43 8. Morphological Analysis of the southeastern smooth sculptured forms of Elimia 51 9. Shell of Melania curvicostata Reeve, 1861 10. Shell of Melania densicostata Reeve, 1861 11. Shell of Goniobasis doolyensis Lea, 1862 12. Shell of Goniobasis elliotti Lea, 1862 13. Shell of Goniobasis etowahensis Lea, 1862 14. Shell of Goniobasis gesneh Lea, 1868 15. Shell of Goniobasis inclinans Lea, 1862 16. Shell of Goniobasis induta Lea, 1862 17. Shell of Goniobasis inosculata Lea, 1862 18. Shell of Melania modesta Lea, 1845 19. Shell of Goniobasis mutabilis Lea, 1862 20. Shell of Goniobasis mutabilis timidus Goodrich, 1942 21. Shell of Goniobasis ucheensis Lea, 1862 xii 22. Shell of Goniobasis viennaensis Lea, 1862 64 23. Aligned mtDNA Sequences of the Cytochrome Oxidase. subunit I of the southeastern Elimia 66 24. Phylogenetic relationships among the smooth sculptured Elimia species in the southeastern river drainages 72 25. Adult shell of Elimia dickinsoni (Lea, 1862), UF 25457 94 26. Adult shell of Elimia species "A", UF 241074, 26.5 mm 99 27. Adult shell of Elimia species "B", UF 222568, 16.3 mm 101 28. Adult shell of Elimia species "C", UF 41451, 18.8 mm 108 xiii Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy ELIMIA CURVICOSTATA SPECIES-COMPLEX WITHIN THE RIVER DRAINAGES OF SOUTHEASTERN UNITED STATES: MORPHOLOGY, DNA AND BIOGEOGRAPHY By Elizabeth Louise Mihalcik December 1998 Chairman: Dr.