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Ann Arbor, MI 48106 SOME ASPECTS OF THE FUNCTIONAL MORPHOLOGY OF THE SHELL OF INFAUNAL BIVALVES (MOLLUSCA) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of the Ohio State University By George Thomas Watters, B.S., M.S. ***** The Ohio State University 1990 Dissertation Committee: Approved by A.S. Gaunt D.H. Stansbery W.C. Sweet Advi ser Department of Zoology B.D. Valentine To my parents and my wife for putting up with it and my grandfather- for starting it all i 1 ACKNOWLEDGMENTS The idea for this study began 14 years ago, and many people during that time have contri buted thoughts and suggesti ons I am indebted to all of these friends, too numerous to mention, but particularly to K. Borror, Dr, R. Fe1tes, W . Kasson, and D r . M . Hoggarth for thei r insights into this research. My committee, Drs. A. Gaunt, D. Stansbery, W. Sweet, and B. Valentine, also offered their suggestions and criticisms. Dr. J. Crites also commented on an ear1y draft. Thi s thesi s mi ght never have been wri tten, and certainly would not have been in this final form, were it not for the generous help of these people. To my wife and all the relatives who gave me support and encouragement, without which I would have given up on this dubious venture, I also offer my thanks and love. Dr. J. Rosewater (U. S. National Museum of Natural History) and Dr. R. Turner (Museum of Comparative Zoology) ki ndly made the col 1ections in thei r care avai1 able to me. This study has been supported by a Sigma Xi Grant-in-Aid of Research, a scholarship from the National Capitol Shell Club, and an Ohio State University Presidential Fellowship. VITA February 28, 1953 ...............Born - Dayton, Ohio 1971 ............................. Graduated, Beavercreek H i gh School, Ohio 1974.. ............................. B.S., University of Miami, Coral Gables, Florida 1973-1977 ........................ Student Co-curator of molluscs, University of Miami, Coral Gables, F 1 or i da 1974-1976 ........................ Enzymology technician, Papanicolaou Cancer Research Institute, Miami, FIori da 1978 ............................. Field entomologist, University of Rhode Island, Kingston, Rhode Island 1979-1981.......................... Micropaleontologist, CLIMAP, University of Rhode Island, Narragansett, Rhode Island 1980 ............................. M.S., University of Rhode Island, Kingston, Rhode Is land 1986.. ............................. Field malacologist, The Nature Conservancy, Columbus, Ohio 1987 ............................. Ecological analyst, Division of Natural Areas and Preserves, Columbus, Ohi o 1987 ............................. Field malacologist, Division of Wildlife, Columbus, Ohio 1988 ............................. Field malacologist, Indiana Department of Natural Resources / U.S. Department of the Interior 1989 ............................. Stream ecologist, Columbia Gas i v PUBLICATIONS Watters, G. T. 1981a. A note on the occurrence of Lithophaga (Leioso lenus) spatiosa (Carpenter, 1857 ) In the shell- plates of Acanthochitona hirudiniformis (Sowerby, 1832). VeTiger 24:77. Watters, G. T. 1981b. Two new species of Acanthochitona from the New World (Polyplacophora: Cryptoplacidae). Nautilus 95:171-177, plates. Watters, G. T. 1983. A new species of Caducifer (Monostiolum) from the western Atlantic (Buccinidae). Nautilus 97:125-128, plates. Watters, G. T. 1986. A survey of the unionid molluscs of the Big Darby Creek System in Ohio. Final Report to The Nature Conservancy, 149 pp., plates, maps. Stansbery, D. H., Stein, C. B., & G. T. Watters. 1986. The distribution and relative abundance of unionid mollusks in the vicinity of Appalachian Power Company’s Clinch River Plant at Carbo, Virginia (Clinch River miles 264- 270). Final Report to American Electric Power Company, 148 pp., plates, maps. Watters, G. T. 1988. The naiad fauna of selected streams in Ohio. I. Stillwater River of Miami River. II. Stream systems of south central Ohio from the LittTe Miami River to the Hocking River, excluding the Scioto River proper. Final Report to the Division of wildlife, Ohio Department of Natural Resources, vi+440 pp. Watters, G. T. 1988. A field-guide to the fresh-water mussels of Ohio. Prepared for the Division of wildlife, Ohio Department of Natural Resources, v+110 pp. Watters, G. T. 1988. A survey of the freshwater mussels of the St. Joseph River system, with emphasis on the federally endangered White Cat’s Paw Pearly Mussel. Final Report to the Division of Fish and Wildlife, Indiana Department of Natural Resources, vi+127 pp. Watters, G. T. & J. Finlay. 1989. A review of the western Atlantic buccinid genera Monostiolum Dali, 1904, and Bailya (Parabailya) new subgenus. Veliger 32:47-59, plates, map. FIELDS OF STUDY Major Field: Zoology v TABLE OF CONTENTS DEDICATION.......................................................... i i ACKNOWLEDGMENTS...................................................i i i VITA i v LIST OF TABLES.................................................... vii LIST OF FIGURES................................................. viii CHAPTER PAGE I. INTRODUCTION.............................................. 18 History of Molluscan Morphometries.............. 23 The Theoretical Morphospace...................... 25 The Paradigm Approach..............................26 Predicted Models of Shell Shape..................27 II. METHODS AND MATERIALS................................. 32 Taxa Used in the Study.............................32 Measurements and Derived Values..................33 III. RESULTS................................................. 43 Comparison of Shell Shapes with Models......... 43 Family Accounts..................................... 48 IV. DISCUSSION..............................................59 Underlying Assumptions and Models............... 59 Evolutionary Considerations...................... 72 SUMMARY..............................................................76 APPENDICES A. Taxa Used In Study..................................... 204 B. Calculated Values of Parameters..................... 218 LIST OF REFERENCES............................................... 232 vi LIST OF TABLES TABLE PAGE 1. Di stri bution of representatives ofthe bivalve families used in study in each of the three morphological phases...................................203 2. Taxa used in study......................................205 3. Calculated values for the taxa used in this study..................................................... 219 vii LIST OF FIGURES FIGURES PAGE 1. Measurements taken from taxa in Appendix A ........ 35 2. Bivalve interpreted as a rectangular solid of length L t width W t and height H ..................... 39 3. Relationship between quasi-streamlining (S) and the offset angle between the line of greatest length and the direction of motion...... 4.1 4. Diagrammatic representation of exchangeable gapage and rocking along a dorso-ventral axis....65 5. Rotation of the valves along the hinge axis (HA)...................................................... 67 6. Rotation of the valves along a dynamic dorso- ventral ax i s .............................................69 7. Anterior and posterior adductor moments for Tresus nuttaJi.......................................... 70 8. Adductor moment lines for Tagelus divisus through entire angle of exchangeable gape rocking movement........................................70 9. Adductor moment lines for Resartia lancaolata through entire angle of exchangeable gape rocking movement........................................71 10. Adductor moment lines for Siliqua patula through entire angle of exchangeable gape rocking movement........................................71 11. Plot of relative permanent gapage