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Mississippian: Osagean) CHONDRICHTHYAN DIVERSITY WITHIN THE BURLINGTON- KEOKUK FISH BED OF SOUTHEAST IOWA AND NORTHWEST ILLINOIS (MISSISSIPPIAN: OSAGEAN) A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science By MATTHEW MICHAEL JAMES HOENIG B.S., Hillsdale College, 2017 2019 Wright State University WRIGHT STATE UNIVERSITY GRADUATE SCHOOL Thursday, September 5th, 2019 I HEREBY RECOMMEND THAT THE THESIS PREPARED UNDER MY SUPERVISION BY Matthew Michael James Hoenig ENTITLED Chondrichthyan diversity within the Burlington-Keokuk Fish Bed of Southeast Iowa and Northwest Illinois (Mississippian: Osagean) BE ACCEPTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF Master of Science Charles N. Ciampaglio, Ph.D. Thesis Director Doyle R. Watts, Ph.D. Chair, Department of Earth & Environmental Sciences Committee on Final Examination David A. Schmidt, Ph.D. Stephen J. Jacquemin, Ph.D. Barry Milligan, Ph.D. Professor and Interim Dean of the Graduate School ABSTRACT Hoenig, Matthew Michael James. M.S. Department of Earth & Environmental Sciences, Wright State University, 2019. Chondrichthyan diversity within the Burlington-Keokuk Fish Bed of Southeast Iowa and Northwest Illinois (Mississippian: Osagean) Chondrichthyan remains occur in abundance within a thin layer of limestone at the top of the Burlington Limestone at the point of the contact with the overlying Keokuk Limestone. This layer of rock, the “Burlington-Keokuk Fish Bed,”1 is stratigraphically consistent and laterally extensive in exposures of the Burlington Limestone near its type section along the Iowa-Illinois border. Deposition of the fish bed occurred on the Burlington Continental Shelf carbonate ramp off the subtropical western coast of Laurussia during the Lower Carboniferous (Late Tournaisian; Osagean) due to a drop in sea level, although the specific mechanism(s) that concentrated the vertebrate fossils remain(s) unknown. Vertebrate remains found within the fish bed include abundant chondrichthyan (Holocephali and Elasmobranchii) teeth, spines, and denticles as well as bony fish and “Acanthodian” (stem chondrichthyan) remains. No studies of these remains or the fish bed itself have been published since 19052; the total chondrichthyan diversity from the fish bed therefore remained relatively unknown until this study. The fish bed 1 The author has chosen to refer to this layer of rock as the “Burlington-Keokuk Fish Bed” instead of the “Burlington-Keokuk Bone Bed” because the first published reference to this stratum describes it as a “Fish” bed instead of a “Bone” bed. 2 This does not include a 2010 Geological Society of America abstract about the fish bed. iii was sampled from two quarries (Nelson Quarry, Mediapolis, IA; Biggsville Quarry, Biggsville, IL). The limestone matrix was dissolved using standard methods for carbonate dissolution, and teeth were identified using original publications from the Burlington and Keokuk Limestones, the Handbook of Paleoichthyology, and appropriate literature. A diverse assemblage of chondrichthyans was recovered that represented at least fifteen genera and twenty three species as well as teeth that did not resemble anything yet described. Additionally, a compilation of all previous studies of the Burlington-Keokuk Fish Bed along with the current study was undertaken in order to create the most complete chondrichthyan faunal list yet reported for the fish bed. These results improve our understanding of chondrichthyan diversity on the Burlington Continental Shelf during the Osagean and will serve as a foundation for future studies of Osagean chondrichthyans both on the Burlington Continental Shelf and elsewhere around the globe. iv TABLE OF CONTENTS Introduction pg. 1 Stratigraphic History of the Burlington Limestone pg. 1 Burlington Limestone Lithology and Stratigraphy pg. 7 Burlington Limestone Age pg. 10 Burlington Limestone Paleogeography pg. 11 Burlington Limestone Depositional Environment pg. 15 Burlington Limestone Sequence Stratigraphy pg. 16 General Paleontology of the Burlington Limestone pg. 18 General Vertebrate Paleontology and History of Study of the Burlington and Keokuk Limestone Vertebrates pg. 21 Formation of the Burlington-Keokuk Fish Bed pg. 29 Hypotheses pg. 34 Methods pg. 36 Data Analysis pg. 41 Results & Discussion pg. 43 Community Diversity pg. 43 Lateral Variation in the Burlington-Keokuk Fish Bed pg. 50 Thin Sections pg. 51 Systematic Paleontology pg. 58 New Species & Taxonomic Revision pg. 87 Avenues for Future Study pg. 87 References pg. 89 v Appendices pg. 97 Appendix I pg. 98 Appendix II pg. 100 Appendix III pg. 108 vi LIST OF FIGURES Figure 1. “Carboniferous Limestones of Iowa” (David Dale Owen, 1852) pg. 3 Figure 2. History of Nomenclature of the Burlington Limestone pg. 8 Figure 3. Isopach Map of Late Tournaisian Strata on the Burlington Shelf pg. 12 Figure 4. Burlington Continental Shelf Paleogeography pg. 13 Figure 5. Burlington Continental Shelf Paleogeography pg. 14 Figure 6. Depositional Facies on the Burlington Continental Shelf pg. 17 Figure 7. Cedar Fork Ooids pg. 18 Figure 8. Relative Sea Levels During the Mississippian pg. 19 Figure 9. Invertebrate Fossils from the Burlington Limestone pg. 20 Figure 10. Crinoid Fossils from the Burlington Limestone pg. 22 Figure 11. Uperocrinus pyriformis. pg. 23 Figure 12. A Sample of the Burlington-Keokuk Fish Bed pg. 24 Figure 13. Conodont Zonations from the Mississippi Valley pg. 25 Figure 14. Bedrock Map & Quarry Locations pg. 37 Figure 15. Nelson Quarry pg. 38 Figure 16. Biggsville Quarry pg. 38 Figure 17. In-situ Burlington-Keokuk Fish Bed pg. 39 Figure 18. Thin Section of the Burlington-Keokuk Fish Bed from Biggsville Quarry pg. 52 vii Figure 19. Thin Section of the Burlington-Keokuk Fish Bed from Biggsville Quarry pg. 52 Figure 20. Thin Section of the Burlington-Keokuk Fish Bed from Biggsville Quarry pg. 53 Figure 21. Thin Section of the Burlington-Keokuk Fish Bed from Nelson Quarry pg. 53 Figure 22. Thin Section of the Burlington-Keokuk Fish Bed from Nelson Quarry pg. 54 Figure 23. Thin Section of the Burlington-Keokuk Fish Bed from Nelson Quarry pg. 54 Figure 24. Thin Section of a “Lesser” Fish Bed from Nelson Quarry pg. 55 Figure 25. Thin Section of a “Lesser” Fish Bed from Nelson Quarry pg. 55 Figure 26. Thin Section of a “Lesser” Fish Bed from Nelson Quarry pg. 56 Figure 27. Thin Section of a “Lesser” Fish Bed from Yarmouth Quarry pg. 56 Figure 28. Thin Section of a “Lesser” Fish Bed from Yarmouth Quarry pg. 57 Figure 29. Thin Section of a “Lesser” Fish Bed from Yarmouth Quarry pg. 57 Figure 30. Chondrichthyan Pharyngeal Denticles pg. 59 Figure 31. Chondrichthyan Pharyngeal Denticles pg. 60 Figure 32. Chondrichthyan Pharyngeal Denticle pg. 61 Figure 33. Chondrichthyan Denticles pg. 62 Figure 34. Helodus sp. pg. 63 viii Figure 35. Deltodus inornatus pg. 64 Figure 36. Venustodus sp. pg. 65 Figure 37. Psammodus sp. pg. 66 Figure 38. Thrinacodus incurvus pg. 67 Figure 39. Cladodus elegans pg. 68 Figure 40. Cladodus bellifer pg. 69 Figure 41. Mesodmodus ornatus pg. 70 Figure 42. Mesodmodus ornatus pg. 71 Figure 43. “Orodus” catanatus pg. 72 Figure 44. “Columnaodus promethii” (New Species) Holotype pg. 73 Figure 45. “Columnaodus promethii” (New Species) Paratypes pg. 74 Figure 46. Orodus mamillaris pg. 75 Figure 47 Orodus varicostatus pg. 76 Figure 48. cf. Orodus elongatus pg. 77 Figure 49. Orodus ramosus pg. 78 Figure 50. Leiodus calcaratus pg. 79 Figure 51. Tanaodus pumilis pg. 80 Figure 52. Tanaodus multiplicatus pg. 81 Figure 53. Tanaodus depressus pg. 81 Figure 54. Tanaodus depressus pg. 81 Figure 55. Chomatodus sp. pg. 82 Figure 56. Antliodus sp. pg. 83 Figure 57. Acanthodii indet. Scales pg. 84 ix Figure 58. Conodont Elements pg. 85 Figure 59. Paleonisciformes indet. Scales pg. 86 Figure 60. Paleonisciformes indet. Teeth pg. 86 Figure 1 (Appendix III) In-situ teeth pg. 108 Figure 2 (Appendix III) In-situ teeth pg. 109 x LIST OF TABLES Table 1. Vertebrate Remains from the Burlington-Keokuk Fish Bed at Nelson and Biggsville Quarries. pg. 43 Table 2. Composite Faunal List for the Burlington-Keokuk Fish Bed. pg. 49 Table 1 (Appendix II) Condrichthyans from the Burlington and/or Keokuk Limestones pg. 100 Table 2 (Appendix II) Condrichthyans from the Burlington and/or Keokuk Limestones that are either synonymous with above species, invalid, inadequately illustrated and described, or taxonomically uncertain pg. 104 xi ACKNOWLEDGEMENTS I would like to thank many people for helping me complete my thesis and coursework at Wright State University. I thank my professors including my academic advisor, Dr. Charles Ciampaglio, the current and previous EES Department Chairs during my time at Wright State, and my committee members, Dr. David Schmidt and Dr. Stephen Jacquemin. Dr. Chad Hammerschmidt and Dr. Silvia Newell provided me with valuable instruction and advice. Dr. Michał Ginter helped recognize a potential new species. Some of the field work was funded by the Richards Field Scholarship Fund. Ryan Shell, David Peterman, and Lauren Fuelling assisted me in the laboratory and field. Lauren deserves special recognition for taking all of the photographs of the thin sections and teeth in the illustrative plates. Preston Meyer, Jerry Malone, and Brad Helmerson from L & W Quarries, Inc., Donald Smith from Cessford Construction Company, and Shawn Duer from Hendricks River Logistics, LLC allowed us to collect samples on their properties. Dr. James E. Day and Dr. Brian Witzke provided valuable advice for my research. The staff at Dunbar Library obtained most of the sources for my research. Dr. J. Nielsen and Dr. R. Phinney helped treat me for a debilitating viral illness during my studies; without their help, I could not have begun, let alone completed, my program of study or thesis. Lastly, I thank my colleagues, friends, family, and church including Jeffrey K., Jeff K., Dr. Larry S., Todd T., Sheldon S., Daniel T., Phillip T., Dr. Anthony S., Dr. Jeffrey V., Cynthia H., Michael, Matt H., Joshua and Kari O., Todd S., Dr. Terry M., Deborah S., Daniel S., Madison E., Shannon C., Wesley A., Megan I., Brian and xii Stephanie K., Caleb and Michelle C., and Michael E.
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