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University Microfilms, Inc., Ann Arbor, Michigan a STUDY of the ANATOMY of the NAIAD This dissertation has been microfilmed exactly as received 69-11,725 YOKLEY, Jr., Paul, 1923- A STUDY OF THE ANATOMY OF THE NAIAD PLEUROBEMA CORDATUM (RAFINESQUE, 1820) (MOLLUSCA: BIVALVIA: UNIONOIDA). The Ohio State University, Ph.D., 1968 Zoology University Microfilms, Inc., Ann Arbor, Michigan A STUDY OF THE ANATOMY OF THE NAIAD PLEUROBMA CORDATUM (RAFINESQUE, 1820) (MOLUJSCA: BIVALVIA: UNIONOIDA) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of the Ohio State University By Paul Yokley, Jr., B.S., M.A ****** The Ohio State University 1968 Approved by jr.s. Adviser Department of Zoology and Entomology ACKNOWLEDGMENTS This study has been made under the supervision of Dr. L. S. Putnam who has been patient when patience counted and prompt when it benefited me most. His help and advice have been most appreciated. To Dr. David H. Stansbery who is a very special friend, thanks is expressed, for the careful way in which he advised, made suggestions, and constructively criticized my work at regular intervals. Dr. John L. Crites has been most helpful in reading the manuscript and providing constructive criticism. His efforts have been greatly appreciated. The cooperative spirit and help provided by Mr. Billy Carroll and Mr. Billy Ison have been greatly appreciated. The effort and skill required in collecting specimens weekly throughout the year would have been inadequate without the help of Mr. Charles Gooch. His diving skill and experience as well as his enthusiasm kept my collecting regular over a two year period. ii Special words of thanks are owed my mother and father who instilled in me the desire to get a good education. Their constant interest and enthusiasm in my work were a continuing help to me. Finally, to my wife, Betty Yokley, much credit is due because she never lost sight of my goal and sacrificed her time to listen and encourage me. In addition, her skill in typing was most important in completing this study. To these and many others who cooperated in so many ways, I shall always be indebted. iii VITA August 3, 1923 Born - Mitchellville, Tennessee 1949 B.S., George Peabody College, Nashville, Tennessee 1950 M.A., George Peabody College, Nashville, Tennessee 1950 Instructor, Florence State College, Florence, Alabama 1960 National Science Foundation Fellow, The Ohio State University, Columbus, Ohio 1961 ..... Assistant Professor, Florence State College, Florence, Alabama PUBLICATIONS "Mussels of the Bear Creek Watershed, Alabama and Mississippi, with a Discussion of the Area Geology." The American Midland Naturalist. 79 (1): 189-196, 1968. "The Mussel Fauna of Duck River in Tennessee, 1965." The American Midland Naturalist. 80 (1): 34-42, 1968. FIELDS OF STUDY Major Field: Zoology Studies in Ornithology. Professor L. S. Putnam Studies in Malacology. Professor D. H. Stansbery iv Major Field: Zoology (continued) Studies in Vertebrate Zoology. Professor J. W. Price Studies in Entomology. Professor D. J. Borror Studies in Botany. Professor C. E. Taft v TABLE OF CONTENTS ACKNOWLEDGMENTS I. INTRODUCTION 1 II. SYSTEMATICS 4 III. DISTRIBUTION 7 1. Range 7 2. Habitat . 8 IV. MATERIALS AND TECHNIQUES 11 V. MORPHOLOGY 17 1. Methods .......... 17 2. Shell 18 3. Mantle ; 24 4. Foot » . ... • 29 5„ Muscular System 31 6. Digestive System 35 7. Circulatory System 56 8. Ctenidial System . 67 9. Nervous System 79 10. Reproductive System .93 11. Excretory System 115 SUMMARY 122 APPENDIX 126 BIBLIOGRAPHY 133 vi LIST OF TABLES Table Page 1. Wilson Dam outlet water temperatures at weekly intervals for 1964 16 2. Wilson Dam outlet water temperatures at weekly- intervals for 1965 126 3. Wilson Dam outlet water temperatures at weekly intervals for 1966 . 127 4. Wilson Dam outlet water temperatures at weekly intervals for 1967 128 5. Wilson Dam outlet water temperatures at weekly intervals for 1968 129 vii LIST OF ILLUSTRATIONS Figure Page 1. Pleurobema cordatum . 5 2. Three of the four closely related forms ... 6 3. Caliper constructed of hardwood . .13 4. Ohio Pigtoe in position to measure length ... 13 5. Ohio Pigtoe in position to measure height ... 14 6. Ohio Pigtoe in position to measure width ... 14 7. Section of mantle margin .23 8. Periostracal gland on margin of mantle.... 23 9. Cardiac muscle viewed longitudinally .... 32 10. Cardiac muscle .32 11. Lateral section of mouth, esophagus and stomach. 37 12. Stomach, lateral section with folds ... .37 13. Median sagittal section of the esophagus and stomach 40 14. Style sac and intestine 41 15. Prointestine and style sac 41 16. Lateral section of the intestinal coils ... 46 17. Transverse section of the mid-intestine ... 46 18. Rectum as it passes through the pericardium . 48 19. Chemosensory tissue on the posterior margin . 48 20. Digestive diverticula .51 21. Transverse section of the secreting acini ... 51 22. Lobule of the digestive diverticulum .... 54 viii Figure Page 23. Transverse section of the ventricle and rectvim . 58 24. Dorsal aorta with lateral branches . 58 25. Anterior aorta with main division 62 26. Pedal sinus in the median section of the foot . 62 27. Transverse section of the dorsal half of a Pigtoe Mussel 64 28. Amoebocytes in the ventricle of the heart ... 64 29. Transverse section of the gill 73 30. Section of a gill through a water tube.... 73 31. Dorsal transverse section of inner laminae... 74 32. Dorsal transverse section where inner gills are connected 74 33. Dorsal transverse section through cloacal chamber . 76 34. Transverse section of outer female gill ... 76 35. Ventral edge of inner gill • 77 36. Transverse section through cerebro-pleural ganglia . 82 37. Multipolar neurons 82 38. Dorsal section of naiad 85 39. Pedal ganglia 85 40. Visceral ganglion 87 41. Sensory tissue . 87 42. Statocyst 92 43* Statocyst 92 44. Median sensory organ of the foot 94 45. Sensory organ of foot 94 46. Testes of Male Pigtoe . .96 ix Figure Page 47. Transverse section of mussel through testes. 96 48. Acini of the testes of a Pigtoe male .... 99 49. Sperm pore and sperm duct . .99 50. Acinus of testes in the spring 103 51. Mature spermatozoa 103 52. Male acinus in summer 105 53. Male acinus in summer 105 54. Male acinus in the fall 106 55. Male acinus in the fall 106 56. Sagittal section through ovaries 108 57. Acini (follicles) of the ovary 108 58. Acini of the ovary of female 112 59. Acini of the ovary 112 60. Acini of ovary from Pigtoe . 113 61. Acinus of ovary 113 62. Several acini of the ovary 114 63. Acinus of ovary 114 64. Renopericardial opening 117 65. Renopericardial opening . 117 66. Afferent arm of the kidney 119 67. External renal orifice 119 68. Profile of Tennessee River showing Dams. .130 69. Depth Profile of Pickwick Reservoir. .... 131 • 70. Diagram of the Digestive System of a Pigtoe. 132 x I. INTRODUCTION The Ohio Pigtoe Mussel, Pleurobema cordatum. in the Family Unionidae, Subfamily Ambleminae, i8 a common ovoviviparous naiad in the Tennessee River. It has had a significant economic history. The Tennessee River was recognized as a source of raw material for buttons as long ago as 1883 when a short-lived pearl button plant was established at Knoxville, Tennessee. In 1914 the Tennessee River furnished 650 tons of mussel shells to the pearl button industry. After the mainstream dams were built, mussel fishermen and many biologists predicted mussels would die out in the river. But in 1945, many species were still alive and the shell harvest was resumed after a lapse of 9 years, rising from 3,700 tons in 1945 to near 10,000 ton3 in 1947- The mainstream reservoirs of the Tennessee River became the most important source of freshwater mussel shells in the United States (T.V.A., 1966). Pearl buttons were gradually replaced by synthetics but the cultured pearl industry created a new demand for the freshwater mussel shells. Again, the Tennessee River naiad shells possessed the most desired characteristics including color, luster and toughness required by this industry. 2 The shell harvest rose annually during the decade following 1945 but in 1956 the harvest was much reduced. After this reduction, the U. S. Fish and Wildlife Service (Scruggs, i960) made a survey of the Pigtoe Mussel which was one of the most desirable commercial species in the Tennessee River. This survey revealed that mussels were being harvested 23 times faster than they were being replaced by younger ones. Also, it showed that the majority of the mussels were older than 12 years. The exact cause for this could not be determined since the life histories of the species of this complex were largely unknown. The mussel harvest has continued to drop annually even though more effort has been expended and more boats and equipment used. Because natural populations of Ohio Pigtoe Mussels have been declining for several years, a renewed study has been made of this species. The emphasis of this study has been on morphology including histological detail and seasonal changes. The study began in the spring of 1964 and continued through the summer of 1968. Included in this study are the seasonal gonadal changes related to oogenesis and spermatogenesis. J The aim of this inquiry was to study as many aspects of the life history, morphology, and ecological requirements as is 3 practicable from samples collected at weekly intervals through­ out several seasonal cycles. A thorough histological study seems to be lacking in this or any closely related species. For this reason, the present work is intended to reveal as much detail of each organ system as possible to provide a foundation for future investigations of this and related species. This study was conducted in the Department of Science, Florence State University, Florence, Alabama.
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