Occurrence and Attributes of Two Echinoderm-Bearing Faunas From

Home , Agaricocrinus, Archaeocidaris, Cleiothyridina, Edrioasteroidea, Listracanthus, Rhombopora

University of Kentucky UKnowledge

Theses and Dissertations--Earth and Environmental Sciences Earth and Environmental Sciences

2018

OCCURRENCE AND ATTRIBUTES OF TWO ECHINODERM- BEARING FAUNAS FROM THE UPPER MISSISSIPPIAN (CHESTERIAN; LOWER SERPUKHOVIAN) RAMEY CREEK MEMBER, SLADE FORMATION, EASTERN KENTUCKY, U.S.A.

Ann Well Harris University of Kentucky, [email protected] Digital Object Identifier: https://doi.org/10.13023/etd.2018.361

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Recommended Citation Harris, Ann Well, "OCCURRENCE AND ATTRIBUTES OF TWO ECHINODERM-BEARING FAUNAS FROM THE UPPER MISSISSIPPIAN (CHESTERIAN; LOWER SERPUKHOVIAN) RAMEY CREEK MEMBER, SLADE FORMATION, EASTERN KENTUCKY, U.S.A." (2018). Theses and Dissertations--Earth and Environmental Sciences. 59. https://uknowledge.uky.edu/ees_etds/59

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The document mentioned above has been reviewed and accepted by the student’s advisor, on behalf of the advisory committee, and by the Director of Graduate Studies (DGS), on behalf of the program; we verify that this is the final, approved version of the student’s thesis including all changes required by the advisory committee. The undersigned agree to abide by the statements above.

Ann Well Harris, Student

Dr. Frank R. Ettensohn, Major Professor

Dr. Edward W. Woolery, Director of Graduate Studies

OCCURRENCE AND ATTRIBUTES OF TWO ECHINODERM-BEARING FAUNAS FROM THE UPPER MISSISSIPPIAN (CHESTERIAN; LOWER SERPUKHOVIAN) RAMEY CREEK MEMBER, SLADE FORMATION, EASTERN KENTUCKY, U.S.A.

______DISSERTATION ______

A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the College of Arts and Sciences at the University of Kentucky

By Ann Well Harris

Lexington, Kentucky

Director: Dr. Frank R. Ettensohn, Professor of Geology

Lexington, Kentucky

2018

ABSTRACT OF DISSERTATION

OCCURRENCE AND ATTRIBUTES OF TWO ECHINODERM-BEARING FAUNAS FROM THE UPPER MISSISSIPPIAN (CHESTERIAN; LOWER SERPUKHOVIAN) RAMEY CREEK MEMBER, SLADE FORMATION, EASTERN KENTUCKY, U.S.A.

Well-preserved echinoderm faunas are rare in the fossil record, and when uncovered, understanding their occurrence can be useful in interpreting other faunas. In this study, two such faunas of the same age from separate localities in the shallow-marine Ramey Creek Member of the Slade Formation in the Upper Mississippian (Chesterian) rocks of eastern Kentucky are examined. Of the more than 5,000 fossil specimens from both localities, only 9–34 percent were echinoderms from 3–5 classes. Nine non-echinoderm (8 invertebrate and one vertebrate) classes occurred at both localities, but of these, bryozoans, brachiopods and sponges dominated. To understand the attributes of both localities (Valley Stone and 213 quarries), the geologic and structural settings, lithofacies and depositional environments, as well as faunal makeup and abundances (diversity, evenness, density), were compared and contrasted. Faunas from the Valley Stone Quarry were located on an uplifted fault block in more shallow, open-marine waters with higher energies. As indicated by four distinct lithofacies, the depositional setting was more extensive and varied with interspersed shoals and basins that could accommodate a greater richness (65 species), even though organism densities and abundance were less. In contrast, fauna from the 213 Quarry were located on a downdropped fault block in a more localized, deeper, storm-shelf setting, characterized by a single lithofacies. Although organism density and abundance were nearly twice as high as that at the Valley Stone Quarry, species richness was lower (45 species), and only one species, a bryozoan, predominated. Overall, echinoderm classes, species and individuals were more abundant at the Valley Stone Quarry, and I suggest that this is related to the shallower and more varied depositional environments that developed in response to presence on the shallow, uplifted fault block. This suggests the importance of regional features like faults in controlling environments and organism distribution through time. Although the faunas were originally collected for their echinoderm-dominated “crinoid gardens,” in fact, echinoderms were in the minority, and bryozoans and brachiopods predominated in the communities. Hence, the communities might better be described as bryozoan “thickets” and brachiopod “pavements.”

KEYWORDS: Late Mississippian, Slade Formation, echinoderms, diversity, richness, depositional environments.

Ann Well Harris

June 28, 2018 Date

OCCURRENCE AND ATTRIBUTES OF TWO ECHINODERM-BEARING FAUNAS FROM THE UPPER MISSISSIPPIAN (CHESTERIAN; LOWER SERPUKHOVIAN) RAMEY CREEK MEMBER, SLADE FORMATION, EASTERN KENTUCKY, U.S.A.

Ann Well Harris

Dr. Frank R. Ettensohn Director of Dissertation

Dr. Edward W. Woolery Director of Graduate Studies

June 28, 2018

ACKNOWLEDGMENTS It is a pleasure to thank many people who made this dissertation possible. Foremost, I want to thank my advisor Dr. Frank R. Ettensohn. It has been an honor to be his Ph.D. student. With his enthusiasm, inspiration, and his efforts to explain things clearly, he helped to make geology enjoyable for me. Throughout my dissertation, he provided encouragement, sound advice, good teaching, and any good ideas. I appreciate all his contributions of time, ideas, to make my Ph.D. experience productive and invaluable. I am also thankful for the excellent example he has provided as a successful geologist and professor. My deepest gratitude is due to the members of the supervisory committee, Dr. Rebecca Freeman, Dr. Alan E. Fryar, Dr. Stephen F. Greb, and Dr. Carol Baskin, without whose knowledge and assistance, this study would not have been successful. My committee members have been an excellent source of advice, feedback, and collaboration. In addition, they have contributed immensely to my personal and professional time at the University of Kentucky. I want to thank my outside examiner, Dr. Timothy Taylor. I want to express my sincere gratitude to Professor Jill Carnahan-Jarvis, Eastern Kentucky University, for her guidance with the statistics, math, graphs, and charts that made this dissertation possible. Her help has been invaluable and has ranged from being my mentor and best friend at a time when I needed someone the most. I would have been lost without her mentorship, encouragement, and friendship. I am thankful for my late master’s thesis advisor, Dr. Robert T. Lierman, Eastern Kentucky University. His encouragement and support throughout the years have been instrumental in my success as a graduate student and adjunct instructor. I will always carry your memory and teaching methods with me. I would like to express my gratitude to Mr. Stephen Moore, Studiospectre, for providing assistance as a paleoartist for the figures and drawings in this dissertation. His efforts are greatly appreciated and made this dissertation possible. I would like to thank Ms. Adrianne Gilley, Department Manager, for her invaluable help and support during my graduate student career at the Department of Earth and Environmental Sciences, University of Kentucky. Lastly, I wish to express my love and gratitude to my family for their understanding and support. Being a first-generation college student, their support has been instrumental in my success. I am grateful for wonderful parents that always believed in me. To my late father, Mr. Billy Wayne Harris, I dedicate this dissertation. Although it has been a few months since you have passed, I carry your life lessons (and my middle name) with me, every day.

Ann Well Harris, Class of 2018 University of Kentucky

iii

TABLE OF CONTENTS

Acknowledgments...... iii

List of Tables ...... vii

List of Figures ...... viii

CHAPTER 1: INTRODUCTION 1.1. Introduction ...... 1 1.2. Purpose of investigation ...... 2 1.3. Hypothesis...... 5 1.4. Study area...... 6 1.5. Methods...... 10 1.5.1. Field and lab methods ...... 10 1.5.2. Statistical methods ...... 13 1.6. Previous studies of Mississippian echinoderm fauna ...... 16

CHAPTER 2: REGIONAL AND LOCAL SETTING 2.1. Regional Structural/Tectonic Framework ...... 18 2.2. Ouachita Orogeny ...... 21 2.3. Local structure influencing the study area ...... 22 2.4. Kentucky River Fault System ...... 23 2.5. Waverly Arch ...... 25 2.6. Paleogeographical/Paleoclimate framework ...... 26

CHAPTER 3: STRATIGRAPHY AND LITHOLOGY 3.1. Stratigraphic framework ...... 29 3.2. History of stratigraphic nomenclature ...... 44 3.3. Slade Formation ...... 46 3.4. Upper part of Slade Formation ...... 46 3.4.1. Poppin Rock Member ...... 46 3.4.2. Maddox Branch Member ...... 47 3.4.3. Ramey Creek Member ...... 47 3.4.4. Tygarts Creek Member ...... 47 3.4.5. Rosslyn Member ...... 48 3.4.6. Holly Fork Member ...... 48 3.4.7. Armstrong Hill Member ...... 48 3.4.8. Cave Branch Bed ...... 48 3.5. Depositional environments ...... 49

CHAPTER 4: FOSSIL-ASSEMBLAGE TAXONOMY 4.1. Systematic paleontology of the new Ophiuroid species ...... 53 4.2. Quarry taxa...... 61 4.2.1. The 213 Quarry ...... 61

4.2.1.1. The 213 Quarry: Top-of-slab ...... 63 4.2.1.2. The 213 Quarry: Bottom-of-slab ...... 65 4.2.2. Valley Stone Quarry ...... 67 4.2.2.1. Valley Stone Quarry: Coarse-grained, calcarenite ...... 70 4.2.2.2. Valley Stone Quarry: Coarse-grained, argillaceous calcarenite .73 4.2.2.3. Valley Stone Quarry: Fine-grained calcarenites and interbedded calcilutite...... 76 4.2.2.4. Valley Stone Quarry: Argillaceous calcilutite ...... 79 4.3. Quarry comparisons ...... 82 4.4. Census populations by class...... 85 4.4.1. Species comparison at 213 and Valley Stone quarries ...... 86

CHAPTER 5: RELATIVE, ORDINAL AND STATISTICAL MEASURES OF ABUNDANCE 5.1. Introduction ...... 94 5.2. Relative abundance ...... 94 5.2.1. Abundance of Demospongea in the 213 and Valley Stone quarries ...... 95 5.2.2. Abundance of Anthozoa in the 213 and Valley Stone quarries ...... 97 5.2.3. Abundance of Scyphozoa (?) in the 213 and Valley Stone quarries ...... 99 5.2.4. Abundance of Pelecypoda in the 213 and Valley Stone quarries ...... 101 5.2.5. Abundance of Polychaeta (?) in the 213 and Valley Stone quarries ...... 103 5.2.6. Abundance of Ostracoda in the 213 and Valley Stone quarries ...... 105 5.2.7. Abundance of Articulata in the 213 and Valley Stone quarries...... 106 5.2.8. Abundance of Inarticulata in the 213 and Valley Stone quarries ...... 109 5.2.9. Abundance of Stenolaemata in the 213 and Valley Stone quarries ...... 111 5.2.10. Abundance of Edrioasteroidea in the 213 and Valley Stone quarries .....114 5.2.11. Abundance of Blastoidea Abundance in the 213 and Valley Stone quarries...... 115 5.2.12. Abundance of Echinoidea in the 213 and Valley Stone quarries ...... 117 5.2.13. Abundance of Ophiuroidea in the 213 and Valley Stone quarries ...... 119 5.2.14. Abundance of Crinoidea in the 213 and Valley Stone quarries ...... 121 5.2.15. Abundance of Chondrichthyes in the 213 and Valley Stone quarries .....125 5.3. 213 and Valley Stone Quarries: comparison of abundances by class ...... 127 5.4. 213 Quarry abundance by class ...... 128 5.4.1. The 213 Quarry: top-of-slab vs. bottom-of-slab abundance by class ...... 128 5.5. Valley Stone Quarry lithofacies abundance by class ...... 130 5.5.1. Lithofacies communities: abundance by class ...... 130 5.6. Ordinal estimates of relative abundance ...... 134 5.6.1. Ordinal estimates of relative abundance for the 213 Quarry ...... 135 5.6.2. Ordinal estimates of relative abundance for the Valley Stone Quarry ...... 139 5.7. Statistical estimates of relative abundance ...... 145 5.7.1. Species Richness ...... 145 5.7.2. Simpson’s Index of Diversity ...... 146 5.7.3. Shannon’s Evenness Index ...... 147 5.7.4. Rank-abundance curves ...... 148

5.7.5. Rank abundance curve for the 213 Quarry: top-of slab vs. bottom-of-slab communities ...... 149 5.7.6. Rank abundance curve for the Valley Stone Quarry: comparison among the four lithofacies communities ...... 150 5.7.7. Rank abundance curve for the 213 vs. Valley Stone quarries ...... 151 5.7.8. Community density ...... 152 5.7.9. Relative frequency ...... 153 5.8. Levels of diversity...... 153 5.8.1. Alpha diversity (α-diversity) ...... 153 5.8.2. Beta diversity (β-diversity) ...... 153 5.8.3. Gamma diversity (γ-diversity) ...... 154 CHAPTER 6: DISCUSSION AND INTERPRETATIONS 6.1. Regional geologic controls ...... 162 6.2. Lithofacies and depositional environments ...... 165 6.2.1. The Valley Stone Quarry lithofacies...... 165 6.2.1.1. Shoal lithofacies ...... 169 6.2.1.2. Shoal-margin lithofacies ...... 172 6.2.1.3. Transitional lithofacies ...... 173 6.2.1.4. Basinal lithofacies ...... 173 6.2.2. The 213 Quarry lithofacies ...... 174 6.3. Taphonomy ...... 180 6.3.1. Biostratinomic indicators for the 213 Quarry communities ...... 181 6.3.2. Biostratinomic indicators for the Valley Stone Quarry communities ...... 195 6.4. Interpretation of relative ordinal and statistical data ...... 212 6.4.1. The 213 Quarry ...... 213 6.4.2. Valley Stone Quarry ...... 215 6.4.3. Locality comparisons ...... 218 6.4.3.1. Depositional environments ...... 218 6.4.3.2. Comparing statistical parameters...... 219 6.4.3.3. Comparing dominant species ...... 222

CHAPTER 7: CONCLUSIONS Conclusions ...... 225 APPENDICES Appendix A – Statistical tables ...... 229

REFERENCES ...... 277

VITA ...... 294

LIST OF TABLES Table 1.1 Area of four sampled lithofacies in the Valley Stone Quarry ...... 11 Table 1.2 Adapted Pearson’s “r” for interpretation of indices...... 15 Table 4.1 Genera/species list for the 213 Quarry ...... 62 Table 4.2 Genera/species for top of slabs, 213 Quarry ...... 63 Table 4.3 Genera/species for bottom of slabs, 213 Quarry ...... 65 Table 4.4 Genera/species list for Valley Stone Quarry ...... 68 Table 4.5 Grabau classification ...... 70 Table 4.6 Genera/species for Valley Stone Quarry, coarse-grained calcarenite lithofacies ...... 70 Table 4.7 Genera/species for Valley Stone Quarry, coarse-grained, argillaceous calcarenite lithofacies ...... 73 Table 4.8 Genera/species for Valley Stone Quarry, fine-grained calcarenites and interbedded calcilutite lithofacies ...... 76 Table 4.9 Genera/species for Valley Quarry, argillaceous calcilutite lithofacies ...... 80 Table 4.10 Comparison of fauna from the 213 and Valley Stone quarries ...... 83 Table 4.11 Quick statistics for the 213 and Valley Stone quarries ...... 85 Table 4.12 Genera/species by assemblage for the 213 Quarry ...... 87 Table 4.13 Genera/species by assemblage for the Valley Stone Quarry ...... 89 Table 4.14 Genera/species by assemblage for the Valley Stone Quarry by lithofacies ....91 Table 5.1 Species abundance descriptors...... 135 Table 5.2 Abundance descriptors for entire 213 Quarry landscape ...... 136 Table 5.3 Abundance descriptors for entire Valley Stone Quarry landscape ...... 141 Table 5.4 Species richness for 213 and Valley Stone quarries ...... 146 Table 5.5 Simpson’s Index of Diversity for 213 and Valley Stone quarries ...... 147 Table 5.6 Shannon’s Evenness Index for 213 and Valley Stone quarries ...... 148 Table 5.7 Community density for 213 and Valley Stone quarries ...... 152 Table 5.8 Species absence/presence for 213 Quarry communities ...... 154 Table 5.9 Species absence/presence for entire Valley Stone Quarry landscape ...... 156 Table 5.10 Alpha, Beta, and Gamma diversities for 213 and Valley Stone landscapes ..159 Table 5.11 Alpha, Beta, and Gamma diversities for top-of-slab and bottom-of-slab communities, 213 Quarry...... 160 Table 5.12 Alpha, Beta, and Gamma diversities for four lithofacies communities, Valley Stone Quarry ...... 160 Table 6.1 Environmental interpretation for lithofacies at 213 and Valley Stone quarries ...... 170 Table 6.2 Three most abundant species for 213 and Valley Stone quarries ...... 181 Table 6.3 Interpreted feeding modes for 213 Quarry ...... 182 Table 6.4 Interpreted feeding tiers for 213 Quarry ...... 185 Table 6.5 Interpreted feeding modes for Valley Stone Quarry ...... 195 Table 6.6 Interpreted feeding tiers for Valley Stone Quarry ...... 198 Table 6.7 Pearson’s “r” interpretation for indicies at the 213 and Valley Stone quarries ...... 214

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LIST OF FIGURES 1.1. Map of study areas ...... 3 1.2. Preservation of echinoid from the Valley Stone Quarry ...... 4 1.3. Map of the 213 Quarry ...... 7 1.4. Map of the Valley Stone Quarry ...... 8 1.5. Outcrop of the Ramey Creek Member ...... 9 1.6. Map of visited outcrop near the Valley Stone Quarry ...... 10 1.7. Photo of slab from the 213 Quarry ...... 13

2.1. Stratigraphic section for Mississippian rocks in central Kentucky ...... 19 2.2. Map of regional depositional environments for Kentucky ...... 22 2.3. Map of structural features in eastern Kentucky ...... 24 2.4. Map of the Mississippian outcrop belt along Cumberland Escarpment ...... 25 2.5. Paleogeographic reconstruction map during Early Carboniferous ...... 27 2.6. Map of the United States during Mississippian time ...... 28

3.1. Exposed units at the Valley Stone Quarry ...... 30 3.2. Stratigraphic section for Mississippian rocks in central Kentucky ...... 31 3.3. Chert replacement at the Valley Stone Quarry ...... 32 3.4. Photo of Ramey Creek Member outcrop ...... 34 3.5. Crossbedding from the coarse-grained calcarenite from Valley Stone Quarry .....35 3.6. Interference ripples from the Valley Stone Quarry...... 35 3.7. Coarse-grained, calcarenite lithofacies from the Valley Stone Quarry ...... 36 3.8. Coarse-grained, argillaceous calcarenite from the Valley Stone Quarry ...... 36 3.9. Fine-grained calcarenite and interbedded calcilutite from the Valley Stone Quarry ...... 37 3.10. Argillaceous calcilutite from the Valley Stone Quarry...... 38 3.11. Stylotitic bedding plane from the Valley Stone Quarry...... 38 3.12. Units exposed at the 213 Quarry ...... 40 3.13. Close-up of the Ramey Creek Member at 213 Quarry ...... 41 3.14. Thin-bedded slab from the 213 Quarry ...... 41 3.15. Preservation and abundance of fossils on the top-of-slab, 213 Quarry ...... 42 3.16. Breccia and deformed limestone from the 213 Quarry ...... 42 3.17. Cross section of top-of-slab and breccia for the 213 Quarry ...... 43 3.18. Graded bedding, gutter cast, and crossbedding from the 213 Quarry ...... 43 3.19. Stratigraphic nomenclature of the Slade and Paragon formations ...... 45 3.20. Generalized Shaw and Irwin model ...... 50 3.21. Stratigraphic section and vertical section from the Valley Stone Quarry ...... 51 4.1. Camera lucida drawing of the arm (oral surface) of Schoenaster n. sp...... 57 4.2. Camera lucida drawing of the oral surface of Schoenaster n. sp...... 58 4.3. Dorsal surface of an arm of Schoenaster n. sp...... 58 4.4. Dorsal view of Schoenaster n. sp...... 59 4.5. Typical ventral surface of Schoenaster n. sp...... 59 4.6. Ventral surface of Schoenaster n. sp...... 60 4.7. Distal arm segment of Schoenaster n. sp...... 60

viii

5.1. Graph of Demospongea abundance in the 213 and Valley Stone quarries ...... 96 5.2. Graph of Demospongea abundance in the four lithofacies at the Valley Stone Quarry ...... 96 5.3. Graph of Anthozoa abundance in the 213 and Valley Stone quarries ...... 97 5.4. Graph of Anthozoa abundance in the 213 Quarry, top-of-slab vs. bottom-of-slab communities ...... 98 5.5. Graph of Anthozoa abundance in the four lithofacies at the Valley Stone Quarry ...... 98 5.6. Graph of Scyphozoa (?) abundance in the 213 and Valley Stone quarries ...... 99 5.7. Graph of Scyphozoa (?) abundance in the 213 Quarry, top-of-slab vs. bottom-of- slab communities ...... 100 5.8. Graph of Scyphozoa (?) abundance in the four lithofacies at the Valley Stone Quarry ...... 100 5.9. Graph of Pelecypoda abundance in the 213 and Valley Stone quarries ...... 101 5.10. Graph Pelecypoda abundance in the 213 Quarry, top-of-slab vs. bottom-of-slab communities ...... 102 5.11. Graph of Pelecypoda abundance in the four lithofacies at the Valley Stone Quarry ...... 102 5.12. Graph of Polychaeta (?) abundance in the 213 and Valley Stone quarries ...... 103 5.13. Graph of Polychaeta (?) abundance in the 213 Quarry, top-of-slab vs. bottom-of- slab communities ...... 104 5.14. Graph of Polychaeta (?) abundance in the four lithofacies at the Valley Stone Quarry ...... 104 5.15. Graph of Ostracoda abundance in the 213 and Valley Stone quarries ...... 105 5.16. Graph of Ostracoda abundance in the 213 Quarry, top-of-slab vs. bottom-of-slab communities ...... 106 5.17. Graph of Articulata abundance in the 213 and Valley Stone quarries ...... 107 5.18. Graph of Articulata abundance in the 213 Quarry, top-of-slab vs. bottom-of-slab communities ...... 108 5.19. Graph of Articulata abundance in the four lithofacies at the Valley Stone Quarry ...... 108 5.20. Graph of Inarticulata abundance in the 213 and Valley Stone quarries ...... 109 5.21. Graph of Inarticulata abundance in the 213 Quarry, top-of-slab vs. bottom-of-slab communities ...... 110 5.22. Graph of Inarticulata abundance in the four lithofacies at the Valley Stone Quarry ...... 110 5.23. Graph of Stenolaemata abundance in the 213 and Valley Stone quarries ...... 112 5.24. Graph of Stenolaemata abundance in the 213 Quarry, top-of-slab vs. bottom-of- slab communities ...... 113 5.25. Graph of Stenolaemata abundance in the four lithofacies at the Valley Stone Quarry ...... 113 5.26. Graph of Edrioasteroidea abundance in the 213 and Valley Stone quarries ...... 114 5.27. Graph of Edrioasteroidea abundance in the four lithofacies at the Valley Stone Quarry ...... 115 5.28. Graph of Blastoidea abundance in the 213 and Valley Stone quarries ...... 116

5.29. Graph of Blastoidea abundance in the 213 Quarry, top-of-slab vs. bottom-of-slab communities ...... 116 5.30. Graph of Blastoidea abundance in the four lithofacies at the Valley Stone Quarry ...... 117 5.31. Graph of Echinoidea abundance in the 213 and Valley Stone quarries ...... 118 5.32. Graph of Echinoidea abundance in the 213 Quarry, top-of-slab vs. bottom-of-slab communities ...... 118 5.33. Graph of Echinoidea abundance in the four lithofacies at the Valley Stone Quarry ...... 119 5.34. Graph of Ophiuroidea abundance in the 213 and Valley Stone quarries ...... 120 5.35. Graph of Ophiuroidea abundance in the four lithofacies at the Valley Stone Quarry ...... 120 5.36. Graph of Crinoidea abundance in the 213 and Valley Stone quarries ...... 122 5.37. Graph of Crinoidea abundance in the 213 Quarry, top-of-slab vs. bottom-of-slab communities ...... 123 5.38. Graph of Crinoidea abundance in the four lithofacies at the Valley Stone Quarry ...... 123 5.39. Graph of Crinoidea by subclass for 213 Quarry locality, top-of-slab and bottom-of- slab communities ...... 124 5.40. Graph of Crinoidea by subclass across the entire Valley Stone Quarry “landscape” ...... 124 5.41. Graph of Chondrichthyes abundance in the 213 and Valley Stone quarries ...... 126 5.42. Graph of Chondrichthyes abundance in the 213 Quarry, top-of-slab vs. bottom-of- slab communities ...... 126 5.43. Graph of Chondrichthyes abundance in the four lithofacies at the Valley Stone Quarry ...... 127 5.44. Graph of comparison abundances for the 213 and Valley Stone quarries ...... 128 5.45. Graph by class in the top-of-slab community at the 213 Quarry locality ...... 129 5.46. Graph by class for the bottom-of-slab community at the 213 Quarry locality ....130 5.47. Graph by class for the coarse-grained, calcarenite community at the Valley Stone Quarry locality ...... 132 5.48. Graph by class for the coarse-grained, argillaceous calcarenite community at the Valley Stone Quarry locality ...... 132 5.49. Graph by class for the fine-grained calcarenite and interbedded calcilutite community at the Valley Stone Quarry locality ...... 133 5.50. Graph by class for the argillaceous calcilutite community at Valley Stone Quarry locality...... 133 5.51. Graph of comparison of top-of-slab and bottom-of-slab communities at 213 Quarry ...... 149 5.52. Graph of comparison of rank-abundance curves for fauna in the four lithofacies communities at the Valley Stone Quarry ...... 150 5.53. Graph of comparison of the entire locality landscape for the 213 vs. Valley Stone quarries ...... 151 6.1. Schematic representation of Upper Slade Formation ...... 164 6.2. Four distinct lithofacies for the Valley Stone Quarry...... 167 6.3. Four distinct lithofacies and interpreted environments ...... 168

6.4. Well preserved crinoid on a stylolite surface ...... 170 6.5. Fossils from on a stylolitic bedding plane from the coarse-grained calcarenite lithofacies (shoal environment) at the Valley Stone Quarry ...... 171 6.6. Aphelocrinus, from coarse-grained calcarenite lithofacies, or shoal environment, in the Valley Stone Quarry...... 172 6.7. Brittle fractures and “slickensides” from the 213 Quarry...... 177 6.8. Rip-up clasts on the bottom-of-slab from the 213 Quarry ...... 177 6.9. Rip-up clasts and load structures from the 213 Quarry ...... 178 6.10. Rip-up clasts, scoured base and storm layer from the 213 Quarry ...... 178 6.11. Ripped up crinoids, scour and disorganized sediment from the 213 Quarry ...... 179 6.12. Taxocrinus, on two slabs from the 213 Quarry...... 179 6.13. Relationships between the top- and bottom-of-slab communities for the 213 Quarry ...... 180 6.14. Schematic reconstruction of the Ramey Creek Member, before-burial top-of-slab community at the 213 Quarry...... 189 6.15. Size-frequency histogram of Taxocrinus, cup width for the 213 Quarry communities...... 192 6.16. Size-frequency histogram of Pentremites, theca height for the 213 Quarry communities...... 192 6.17. Photo of slab cut through a crossbedded calcarenite from the shoal environment ...... 202 6.18. Blastoid preserved as thecae...... 203 6.19. Size-frequency histogram of Phanocrinus, cup width from the Valley Stone Quarry shoal community ...... 203 6.20. Schematic reconstruction of Ramey Creek Member environmental continuum .204 6.21. Bedding plane from the fine-grained calcarenite and interbedded calcilutite lithofacies (shoal-margin environment) ...... 206 6.22. Size-frequency histogram of Phanocrinus, cup width for the Valley Stone Quarry shoal-margin community ...... 206 6.23. Size-frequency histogram of Phanocrinus, cup width for the Valley Stone Quarry transitional community ...... 208 6.24. Size-frequency histogram of Phanocrinus, cup width for the Valley Stone Quarry basinal community ...... 209 6.25. Preservation of Aphelocrinus, preserved in argillaceous calcilutite from the basinal environment ...... 211 6.26. Agassizocrinus from the argillaceous calcilutite lithofacies ...... 211 6.27. Thin graded beds from the argillaceous calcilutite lithofacies ...... 212 6.28. Phanocrinus, with two extremely thick arms protecting the more delicate inner arms ...... 224

CHAPTER 1: INTRODUCTION

1.1. Introduction

Today, the echinoderms are among the most numerous and diverse of marine invertebrates (Waggoner, 1999; Brusca and Brusca, 2003), and in the Carboniferous, they were just as abundant (Ettensohn et al., 2003). In particular, during the Mississippian time, echinoderms underwent significant evolution and dramatically increased in abundance and diversity (Ettensohn et al., 2003; Zamora and Rahman, 2017). In fact, one echinoderm group became so abundant during that the Mississippian Period is frequently referred to as the “Age of the Crinoids” (Lane, 1972; Levin, 1999; Kammer and Ausich,

2006). This project examines two Late Mississippian (Chesterian, early Serpukhovian) echinoderm communities of about 328 million years in age (Davydov et al., 2012), one from northeastern and one from east-central Kentucky (Figure 1.1), to ascertain their paleoecology and to examine the differences between the two communities. The paleoecology of echinoderm communities has been studied from the Ordovician

(Whittington, 1963; Sprinkle and Guensburg, 1995; Ausich, 1999a; Brett, 1999a; Brett et al., 2008; Botting et al., 2013; Brower, 2011, 2013; Reich et al., 2017; Milam, et al.,

2017; Cole et al., 2018); Silurian (Taylor and Brett, 1996, 1999; Hess, 1999a; Sumrall et al., 2013); Devonian (Brett et al., 1999b; Hess, 1999b; Thompson et al., 2013);

Mississippian (Chesnut and Ettensohn, 1988; Ettensohn and Chesnut, 1989; Ausich and

Meyer, 1992, 1994; Ausich, 1999b, c); Pennsylvanian (Ausich, 1999d; Sumrall et al.,

2006), and Permian (Hess, 1999c; Webster and Lane, 2007) rocks. These previous echinoderm community studies have largely dealt with systematics rather than organism interrelationships. Moreover, when organism relationships are discussed, the

relationships are typically among echinoderms (e.g., Brower, 2013). In addition, most of these studies, only emphasize the analysis of one community. In addition, the only

Chesterian echinoderm community studies among all the communities studies examined are those by Chesnut and Ettensohn (1988) and Ettensohn and Chesnut (1989), and these studies are from younger Chesterian rocks (upper Hombergian) than those in this project.

In this project, I analyzed two different echinoderm-dominated communities along with their non-echinoderm community members from the same lower Hombergian rocks. This study is the first in-depth and extensive examination of the interrelationships among different organisms in a Late Misssissippian community.

1.2. Purpose of Investigation

The purpose of this project is to identify and analyze echinoderm faunas along with other community members to characterize organism interactions, facies, and possibly even regional structural influences, on the assemblages. The specimens were collected from the Ramey Creek Member of the upper Slade Formation of the mid-

Chesterian (early Serpukhovian) Series from northeastern and east-central Kentucky.

Limestone slabs were collected from two different echinoderm communities in quarries separated by 80 km (50 mi) (Figure 1.1).

Figure 1.1. The two black asterisks represent the approximate locations of the two faunas collected in Carter and Powell counties, Kentucky. The green outcrop belt in eastern Kentucky represents lower to middle Mississippian rocks. The turquoise outcrop belt is the Slade Formation.

The Ramey Creek Member represents a shallow, open-marine setting and contains the most diverse and populous fauna of any Slade member (Ettensohn et al.,

2002). Among the most common faunal elements in the Ramey Creek Member are well- preserved echinoderms. Because echinoderm skeletons are composed of many isolated plates and disaggregate rapidly after death articulated specimens are rare (Fenton and

Fenton, 1958; Meyer and Meyer, 1986; Allison, 1990; Baumiller and Ausich, 1992;

Ausich and Baumiller, 1998; Baumiller, 2003; Gahn and Baumiller, 2005; Lin et al.,

2008; Gorzelak and Salamon, 2013; Deline and Thomka, 2017). However, in the two

study areas, nearly all the organisms are complete and fully articulated, providing an unusual opportunity to examine the echinoderms and other community members individually as species and as members of communities (Figure 1.2). This is the first paleoecology study of echinoderm faunas from the lower Hombergian Substage of the

Chester (early Serpukhovian).

Figure 1.2. This echinoid specimen from the Valley Stone Quarry is complete and fully articulated, providing an unusual opportunity to examine its features. The specimen is largely replaced by red chert.

1.3. Hypothesis

Echinoderms have been collected and studied from the Slade Formation during the last 30‒40 years (Ettensohn and Chesnut, 1985; Chesnut and Ettensohn, 1988;

Ettensohn et al., 2009). Two echinoderms faunas were collected from the Slade

Formation in the 1970s and were available for study. Although the two collections are from the same member, of the same age, and relatively close together (83 km [50 mi]), the lithology and faunal composition of the two communities are strikingly different. I hypothesize that the differences between the two echinoderm faunas and their occurrences may reflect regional and local structural factors that controlled the depositional settings. If I can show that the nature of the communities is virtually the same, then I could falsify the hypothesis. On the other hand, if I can prove that the communities are different and can convincingly relate these differences to structural control, then I have supported the hypothesis, at least in principle. I propose to do the following: characterize the fauna, diversity, life modes, and abundances from each community descriptively and statistically so that they can be compared; characterize the depositional environment(s) in which each community occurs by examining lithology, sedimentary structures and stratigraphic setting; and characterize the structural setting of each community. I will then integrate the results for a comprehensive study of the paleoecology of the communities. This will be the first detailed study of the paleoecology of these faunas from the Upper Mississippian Ramey Creek Member of the Slade

Formation.

1.4. Study Area

The study area includes a location known as the 213 Quarry in Powell County,

Kentucky (Figure 1.3). Ninety slabs of rock, representing a single bed, were collected from an area of approximately 35 ft² (3.3 m²) along with many loose specimens. The second study area, the Olive Hill Ken-Mor Quarry locality, now known as the Valley

Stone Quarry, is located approximately 80 km (50 mi) to the northeast of the 213 Quarry, in Carter County, Kentucky (Figure 1.1). Specimens from the Valley Stone Quarry are included on 136 slabs and include many loose specimens from multiple beds. Overall, the area in this quarry that exhibits the echinoderm-bearing facies may have been more than

0.17 km² (0.07 mi²) (Figure 1.4). Since the Valley Stone Quarry is no longer accessible, I visited an outcrop adjacent to the Valley Stone Quarry to examine lithologies and features of the Tygarts Creek Member and Ramey Creek Member (Figure 1.5). Figure 1.6 shows the location of the visited adjacent outcrop (lower red star) with all characteristic

Ramey Creek Member lithofacies.

Figure 1.3. Location of the 213 Quarry, Powell County, Kentucky, east of Kentucky State Highway 213.

Figure 1.4. Location of the Valley Stone Quarry between I-64 and Kentucky State Highway 182, showing the large extent of the quarry across which specimens were collected.

Figure 1.5. An outcrop of the Ramey Creek Member adjacent to the Valley Stone Quarry (see Figure 1.6), located along U.S. Highway 60, Carter County, KY, used to examine lithologies and sedimentary features.

Figure 1.6. Map showing the location of the outcrop in Figure 1.5 (lower red star) with all characteristic Ramey Creek lithofacies (modified from Google Maps). The Valley Stone Quarry is no longer accessible.

1.5. Methods

1.5.1. Field and Lab Methods

Specimens in this study were collected from the two quarries shown in Figure

1.1. Most of the specimens are on small to large slabs that include lithologies and sedimentary structures, as well as examples of all echinoderm and non-echinoderm fauna.

At the 213 Quarry, the entire community was collected from one lithology (facies) in a corner of the quarry, the area of the slabs (35 ft²; 3.25 m2) represents the approximate areal extent of the former echinoderm community. At the Valley Stone Quarry, however,

specimens occurred in four different lithologies (lithofacies) across a broad part of the quarry, and collecting all specimens was impossible. After each blasting event, all accessible specimens were collected, but it was clear at the time that many specimens could not be collected because of slab size. In an attempt to provide some type of normalization among the lithofacies at each quarry, the approximate areas of the collected slabs in each Valley Stone lithofacies were measured so that some measurement of relative specimen density per unit area could be provided. The approximate collected areas for each lithofacies at the Valley Stone Quarry are shown in Table 1.1.

Table 1.1 Approximate areas of the four sampled lithofacies present in the Valley Stone Quarry.

Approximate area of the four sampled lithofacies present in the Valley Stone Quarry Environment Number of Area in by lithology Slabs ft2 (m2) Coarse- grained calcarenite 21.47 60 (1.99) Coarse- grained, argillaceous 9.66 calcarenites 25 (0.89) Fine-grained calcarenites and interbedded 6.22 calcilutites 19 (0.57) Argillaceous 6.41 calcilutites 32 (0.59) 43.76 Total 136 (4.06)

Echinoderms were identified using standard references and taxonomic procedures

(e.g., McFarlan, 1942; Kirk, 1939, 1940, 1944a, b; Horowitz, 1965; Perry and Horowitz,

1963; Utgaard and Perry, 1969; Chesnut and Ettensohn, 1988; Kammer and Springer,

2008). Beerbower (1968) suggested that most species and genera occur within a limited range of sedimentary environments. Therefore, the rocks and unit sequences in which the echinoderms occur were carefully examined for environmental indicators.

Specimens were carefully unwrapped, washed and cleaned with toothbrushes and dental picks, and allowed to dry. Many of the specimens were covered with a thin parting of shale (Figure 1.7) that had to be carefully removed for identification of specimens, but the presence of the shale itself was an indicator of an important environmental event, and so one slab with a shale covering was preserved (Figure 1.7). Loose specimens were collected from the surface of each slab in order to include them in the study data.

Important and representative specimens were photographed or drawn with camera lucida.

Each rock slab/specimen was labeled with a number and stored for identification.

Fossils were identified to the species or genus level, using an AmScope trinocular

LED stereo microscope, a binocular microscope or hand lens. Identifying some individuals to the species level was not possible owing to their poor preservation, rendering them unidentifiable even to the genus level at times. However, most community members were identified to at least the genus level, using several publications to help with identification (e.g., Worthen, 1883, 1870, 1875, 1890; McFarlan, 1942; Kirk,

1939, 1940, 1944a, b; Horowitz, 1965; Perry and Horowitz, 1963; Utgaard and Perry,

1969; Strimple, 1975; Chesnut and Ettensohn, 1988; Kammer and Springer, 2008).

Specimen counts of each species were tallied for each rock slab and recorded in lab notes.

All echinoderm and non-echinoderm species were tallied, and these tallies were recorded in a Microsoft Excel spreadsheet for data analyses. Sedimentologic and stratigraphic features were also noted on each slab, and the stratigraphy of the original localities was logged.

Figure 1.7. Slab from the 213 Quarry showing bryozoan, crinoid and brachiopod specimens in original state, covered by a thin veneer of blue-gray shale.

1.5.2. Statistical Methods

To compare the fossil organisms and communities at each quarry, I analyzed the census-population data (Appendix A; Tables TA1 – TA12) for richness, diversity and evenness using several statistical methods. All statistical analyses were performed using

Microsoft Excel. Data for comparing richness, diversity, and evenness were analyzed using census-population counts from the top and bottom-of-slabs from the 213 Quarry, as well as for collections from the four different lithofacies at the Valley Stone Quarry.

Overall, it is important to understand how the diversity compares across various

communities, looking at diversity as the number of different kinds of organisms in each community and a community as an assemblage of organisms inhabiting a specific area

(Johnson, 1964). One of the measures of diversity is species richness (S), which is defined as the total number of species or taxa (S = number of species or taxa) (Fairbridge and Jablonski, 1979; Stanton, 1979; Stirling and Wilsey, 2001). Species richness reflects species presence, but does not factor in the abundance or absence of species.

In developing these comparisons, understanding how diverse a community is, or the degree of diversity, can be extremely useful in making comparisons. This aspect of diversity can be measured using Simpson’s Index D (Simpson, 1949) for an entire community. D = (∑ n*(n-1)) / (N*(N-1)), where n = total number of individuals of a species and N = sum of individuals of all species. Simpson’s Index of Diversity is 1 – D, and the result will be a number between 0 and 1. Numbers close to 0 indicate a lack of diversity and numbers close to 1 indicate strong diversity.

Another measure of diversity is evenness (EH), which describes how uniformly abundant taxa are in a community. Shannon’s Diversity Index H was used to calculate

Shannon’s Evenness Index EH (Shannon and Weaver, 1949). H = ∑ - (P * ln P), where P

= proportion (n/N) and lnP is the natural logarithm of P. Shannon’s Evenness Index EH =

H / ln (S), and the result is a number between 0 and 1. Numbers close to 0 indicate little to no evenness (one species is dominating the community), and numbers close to 1 indicate strong evenness (multiple species predominate). It is important to note if no species were present in the community, then the proportion, P, would be equal to 0, and the ln (0) is undefined mathematically (Appendix A; Tables TA1 – TA12). I have

adjusted the number of species in these calculations to include only those present in each community respectively.

An adaptation of the Pearson correlation coefficient, also referred to as

Pearson’s “r” (Franzblau, 1962; Hinkle et al., 1988), is a statistical measurement, ranging from 0 to ±1, which provides an indication of how strong or weak a linear relationship between two variables. I used Pearson’s “r” as a “rule of thumb” to interpret both Simpson’s Index of Diversity and Shannon’s Evenness Index. In both calculations,

Simpson’s Index of Diversity and Shannon’s Evenness Index use the laws of probability and interpretations resulting from them. The higher the value, the more likely the outcome is to occur. Because no interpretative values were available to reflect the frequently used descriptive terms, weak, moderate and strong, I developed an adaptation of Pearson’s “r”, for comparative purposes (Harris et al., 2018). The relationship between these descriptive terms and the probability measurements for each diversity measure using Pearson’s “r” are shown in Table 1.2 and this relationship is used as a general “rule of thumb.” The related calculations will be discussed later in Chapter 6 and Appendix A;

Tables TA1 – TA12.

Table 1.2 Adapted Pearson’s “r” interpretation for this study.

Adapted Pearson’s “r” for Interpretation of Indices Simpson’s Diversity and Interpretation of Index Shannon’s Evenness Index No statistical significance for Diversity or 0 – 0.20 Evenness 0.20 – 0.40 Weak Diversity or Evenness 0.40 – 0.60 Moderate Diversity or Evenness 0.60 – 0.80 Somewhat Strong Diversity or Evenness 0.80 – 1.00 Strong Diversity or Evenness

Other statistical analyses used for understanding community diversity were density and relative frequency. Density for the community is defined as ∑ n / Area, where n = total number of individuals of all species. However, density for each individual species is n / Area. The number of individuals per square meter was determined, providing insight into relative abundance. The relative frequency of an individual species is the number of individuals compared to the total number of all individuals in its environment. The relative frequency represents the probability that a randomly selected sample will be from that particular species. The relative frequency = n/N, where n = total number of individuals of a species, and N = sum of individuals of all species.

1.6. Previous Studies of Mississippian Echinoderm Fauna

The Mississippian rock record has been investigated over the past few decades by many researchers who have focused on echinoderms, and more specifically, on crinoid paleoecology. In fact, the Mississippian has been referred to as the “Age of the Crinoids”

(Kammer and Ausich, 2006; Ettensohn et al., 2007; Ettensohn et al., 2009). The majority of echinoderm studies, however, have been conducted in Kinderhookian and Osagean rocks from Kentucky, Indiana and Tennessee (Van Sant and Lane, 1964; Lane and

Dubar, 1983; Kammer, 1984; Meyer et al., 1989; Feldman, 1989; Ausich and Meyer,

1992, 1994; Kammer and Ausich, 1996; Meyer and Ausich, 1997; Ausich et al., 1997,

2000; Lee et al., 2005; Rhenberg et al., 2016). Moreover, only a few authors have conducted comprehensive studies of crinoid faunas from the Lower Mississippian section in other states and provinces, including Utah (Ausich, 2003), Montana (Laudon and

Severson, 1953) and Alberta, Canada (Laudon et al., 1952).

A major change in echinoderm faunas involving the loss of large, many-plated camerate crinoids occurred in Meramecian time (Ettensohn et al., 2009), but only a few studies of echinoderms are available for this interval. Laudon (1957) and Maples et al.

(1995) described some of the above changes, and a study by Cook (2010) focused on the systematics and evolutionary paleoecology of crinoids from the Middle Mississippian St.

Louis Limestone (Mississippian, Meramecian) of the Illinois Basin.

Only a few studies have focused on Late Mississippian (Chesterian) echinoderms, and these include research from Alabama (Horowitz and Waters, 1972; Burdick and

Strimple, 1982), Arkansas (Burdick and Strimple, 1973), Kentucky (Ettensohn and

Chesnut, 1979; Chesnut and Ettensohn, 1988; Ettensohn et al., 2009), Tennessee

(Strimple and Coney, 1982; Knox and Kendrick, 1987), and West Virginia (Kammer and

Springer, 2008). Of the above research, none have focused on Hombergian (Middle

Chesterian) echinoderms like those from the Ramey Creek Member of the Slade

Formation. The only closely related echinoderm community research is from the Wymps

Gap Member of the Greenbriar Formation (Kammer and Springer, 2008), which is slightly older than the studied rocks in this project, and from younger parts (Poppin Rock

Member) of the Slade Formation in south-central and southeastern Kentucky (Ettensohn and Chesnut, 1979; Chesnut and Ettensohn, 1988) and equivalent rocks in Indiana

(Horowitz, 1965). Ettensohn et al. (2007, 2009) have detailed the biostratigraphy of all significant Mississippian echinoderms from the Appalachian Basin. Of these studies, none of have focused on Hombergian (Middle Chester) echinoderms like those from the

Ramey Creek Member of the Slade Formation.

CHAPTER 2: REGIONAL AND LOCAL SETTING

2.1. Regional Structural/Tectonic Framework

The Mississippian System of the Central Appalachian Basin is part of a foreland- basin succession (Ettensohn et al., 2002), which exhibits a three-part stratigraphic sequence composed of Lower and Middle Mississippian (Tournaisian–early Viséan;

Kinderhookian–Osagean) clastic sediments (Sunbury and Borden formations), Middle and Upper Mississippian (middle–late Viséan; Meramecian–early Chesterian) carbonates

(Slade Formation), and uppermost Mississippian (latest Viséan–Serpukhovian; late

Chesterian) (Figure 2.1) clastic sediments (Paragon Formation), which are respectively interpreted to represent post-Acadian clastic influx, widespread carbonate deposition accompanying tectonic quiescence, and renewed clastic influx marking inception of the

Alleghanian Orogeny (Rodgers, 1970; Perry, 1978; Chesnut, 1991).

The Mississippian rocks in the Appalachian Basin are some of the most interesting rocks in Kentucky because of their diverse nature. The Middle and Upper

Mississippian (Meramecian and Chesterian) carbonates of east-central and northeastern

Kentucky have been extensively studied for decades (e.g., Dever, 1980, 1999; Ettensohn,

1980, 1991, 1992; Ettensohn et al., 1984, 1988, 2002, 2009; Sable and Dever, 1990; Al-

Tawil and Read, 2003; Wilhelm, 2008; Zeng et al., 2013), and are interpreted regionally to represent deposition across a broad, ramp that dipped gently eastward into the

Appalachian foreland basin.

Figure 2.1. The regional stratigraphy of Upper Devonian, Mississippian and Lower Pennsylvanian rocks in central Kentucky. The red arrows show the stratigraphic location of the Ramey Creek Member, the unit being studied for this project (adapted from Ettensohn, 2009).

The Mississippian section in the Appalachian Basin occupies a flexural foreland basin that was influenced by three main orogenies: the Acadian/Neoacadian, Alleghanian, and Ouachita, and varies in thickness from approximately 30 meters to 1,900 meters

(Ettensohn, 2009).

The Acadian/Neoacadian Orogeny represents a north-to-south, zipper-like, transpressive collision between the southeastern margin of Laurussia and Avalonian terranes, which began in Early Devonian time and persisted until the end of Mississippian time (Ettensohn, 2008, 2009; Lierman et al., 2011). The most active deformational parts of the orogeny, however, occurred during Middle and Late Devonian time, and by the

Late Mississippian (Chesterian), tectonic relaxation ensued throughout the Appalachian

Basin. One of the major aspects of relaxation was the deposition of a thin, widespread blanket of carbonates across the basin area, of which the Slade Formation was a part

(Ettensohn et al., 1984; Ettensohn, 1994). Older concepts of the Acadian orogeny include what we call today the Neoacadian orogeny (Robinson et al., 1998). However, some authors (e.g., Hatcher et al., 2005) have interpreted Mississippian events in the southern and central Appalachians as reflective of forces generated by Neoacadian orogeny, and hence, the Slade carbonates represent one part of the relaxation phase of the Neoacadian orogeny (Ettensohn, 1998, 2008).

The Appalachian area at this time was part of the Acadian/Neoacadian foreland basin and was covered by a shallow inland sea. A belt of Neoacadian Highlands bordered the continent and separated the inland sea from waters of the Rheic Ocean, but these highlands had little or no influence on Mississippian environments. Gondwana eventually closed the Rheic Ocean via convergence with Laurussia in Pennsylvanian-Permian time.

The closing of the Rheic Ocean, which separated Laurussia and Gondwana, affected the oceanic and atmospheric circulation patterns (Smith and Read, 2000), and in part, contributed to the continental glaciations that formed in the southern hemisphere on

Gondwana, thereby generating sea-level fluctuations as the glaciers repeatedly formed and melted.

2.2. Ouachita Orogeny

The Ouachita Orogeny along the southern margin of Laurussia involved the collision of South America parts of Gondwana with Laurussia. The orogeny apparently began in latest Devonian time and was ongoing throughout Mississippian and

Pennsylvanian time. Ettensohn (1993) has suggested that this orogeny in part may have controlled the distribution of carbonates in the study area. The Ouachita Orogeny (Figure

2.2) was the final phase of the collision between Laurussia and Gondwana and probably controlled reactivation of structures related to the Kentucky River Fault System and

Waverly Arch through bulge moveout (Zeng et al., 2013).

Figure 2.2. Map showing regional depositional environments (shallow marine) for Kentucky, relative to the locations of the paleoequator, Ouachita foredeep and the Cincinnati Arch (modified from Smith and Read, 2000).

2.3. Local Structure Influencing the Study Area

Northeastern and east-central Kentucky are underlain by three structures which were apparently active during Carboniferous time (Ettensohn, 1992): two broad, east- west basement fault systems with surface expressions in the Kentucky River and Irvine-

Paint Creek fault systems and a north-south-trending uplift named the Waverly Arch by

Woodward (1961) (Figure 2.3).

All of these structures were apparently active during the Early Paleozoic time

(Woodward, 1961) and appear to have left northeastern Kentucky predominately positive during Carboniferous time (Dever, 1977, 1980; Ettensohn, 1977, 1980, 1981b). Much of

the structural movement probably reflects bulge reactivation of basement structures during Ouachita tectonism and Acadian relaxation (Ettensohn, 2009; Zeng et al., 2013).

2.4. Kentucky River Fault System

The Rome Trough Fault System is a Late Precambrian–Early Cambrian continental rift zone that developed during the late Iapetan rifting event (Zeng et al.,

2013; Thomas, 1991; Ettensohn, 1992), and it underlies a major part of the study area.

The trough is a subsurface, graben-like structure that extends from south-central

Kentucky through central and eastern Kentucky into West Virginia (Thomas, 1991). In

Kentucky, three fault systems bound the Rome Trough: the Kentucky River Fault

System, the Lexington Fault System, and the Rockcastle River-Warfield Fault System

(Goodman, 1992; Drahovzal and Noger, 1995; Zeng et al., 2013).

The Kentucky River Fault System, one of the major structural features in

Kentucky, is a narrow band of surface and subsurface normal faults and grabens trending east-northeast from Casey County to the Kentucky-West Virginia border (Figure 2.3).

The faults are at the surface from Casey County to Montgomery County, but reactivated subsurface equivalents are present from Montgomery County eastward to West Virginia

(Figure 2.3). The displacement is to the southeast, and the total throw is as much as 600 feet (Black and Haney, 1975).

Figure 2.3. Structural features (Kentucky River and Irvine-Paint Creek fault systems and Waverly Arch) in eastern Kentucky showing the location of the study area in Carter and Powell counties (figure modified after Harris et al., 2003).

The Kentucky River Fault System forms a series of down-to-the-south normal faults, which mark the northern margin of the Precambrian Rome Trough (Ammerman and Keller, 1979). Dever (1977) and Ettensohn (1977) have shown that this fault system was periodically reactivated as subsurface growth faults during the deposition of the

Mississippian carbonates of the study area. The Valley Stone Quarry in Carter County is on the northern uplifted fault block of this system, whereas the 213 Quarry locality is located south of the faults on a downdropped block (Figure 2.4). In general, carbonates deposited in the north are coarser-grained and reflect higher energy environments

(Ettensohn, 1977, 1980, 1981a, b). A major purpose of this study is to determine if the

fault system had any influence on the nature and development of the studied echinoderm communities.

Figure 2.4. Map showing the Mississippian outcrop belt along the Cumberland Escarpment. The Valley Stone Quarry is located on the northern uplifted block of the Kentucky River Fault System, whereas the 213 Quarry is located on the southern downdropped block. Other regional structures are also shown (adapted from Ettensohn, 1977).

2.5. Waverly Arch

The Waverly Arch is another prominent structural feature that probably influenced Chesterian depositional environments (Ettensohn, 1977, 1980, 1986).

Woodward (1961) first described the Waverly Arch as a broad, concealed arch east of the larger Cincinnati Arch, extending from central Ohio to east-central Kentucky, and both

Pashin and Ettensohn (1987) and Ettensohn (1992) have suggested that the arch reflects uplift on a basement fault. The influence of the Waverly Arch on the deposition of parts 25

of the Slade limestone in northeastern Kentucky was first recognized by Dever (1973), based on thinning and peculiar subaerial diagenesis of lower Slade members. The

Waverly Arch coincides with the Mississippian outcrop belt for nearly half its length in east-central Kentucky and appears to have significantly influenced Carboniferous sedimentation in the study area (Ettensohn, 1992); its influence was apparently far more pronounced on the northern uplifted block of the Kentucky River Fault Zone (Ettensohn,

1981a, b).

The Waverly Arch and Kentucky River Fault System influenced Cambro–

Ordovician and Carboniferous deposition (Woodward, 1961; Ettensohn and Dever, 1975;

Dever, 1977; Ettensohn, 1977). Stratigraphic relationships between the linear Carter

Caves Sandstone and underlying units indicate that uplift of the Waverly Arch was probably concurrent with middle Chesterian activity along the Kentucky River Fault

Zone (Ettensohn and Peppers, 1979).

The Waverly and Cincinnati arches were reactivated periodically throughout

Phanerozoic time and disrupted regional depositional processes (Zeng et al., 2013).

Reactivation of these structures was likely a major factor controlling the distribution of sedimentary facies (Ettensohn, 1980; Dever, 1999; Root and Onasch, 1999; Ettensohn,

2008).

2.6. Paleogeographic/Paleoclimate Framework

During Mississippian time, the North America craton straddled the paleoequator so that much of the United States was situated in the trade-wind belt, 5–30° south of the paleoequator (Scotese, 2003; Ettensohn, 2001) (Figure 2.5). The Euramerica

supercontinent, or Laurussia (Figure 2.5), formed with the collision of Baltica and

Laurentia during the Silurian Caledonian orogeny and was largely an equatorial continent. The study area was located approximately 25° south of the paleoequator

Figure 2.5. Paleogeographic reconstruction showing approximate position Euramerica (Laurussia) and Gondwana supercontinents. During Early Carboniferous (Mississippian) time, the study area was located about 20° south of the paleoequator (adapted from Scotese, 2003). The red dot shows the approximate position of the study area in the Appalachian basin. in the subtropical, trade-wind belt (Figure 2.5). The location of the exact study area is shown above with the red dot. The study area was covered by a shallow epicontinental sea (Figure 2.2).

During most of this time, southern parts of the Laurussian continent, including

Kentucky, were covered by warm, shallow, well-lit, inland seas on the margin of the

Appalachian foreland basin. Shallow seas extended inland from the deep ocean at the edge of the continental plate (Figure 2.6). Figure 2.5 and 2.6 shows the approximate location of Kentucky in relation to the paleoequator.

Figure 2.6. Map showing the approximate position of the United States during Mississippian time. Warm, shallow, well-lit, inland seas covered Kentucky on the margin of the Appalachian foreland basin (red dashed line) (adapted from Illinois State Geological Survey, 2018).

A belt of mountains arose during the Neoacadian orogeny along the eastern margin of Laurussia (Figure 2.6). By Late Mississippian time, however, sea level had dropped (Ettensohn, 2009), carbonate deposition waned and the shallow seas were inundated by clastic sediment eroded from rising Neoacadian highlands to the east. The

Appalachian Basin also experienced infilling, as the tectonic highlands rebounded after unloading during late Chesterian time (Ettensohn et al., 2002).

CHAPTER 3: STRATIGRAPHY AND LITHOLOGY

3.1. Stratigraphic Framework

Stratigraphically, the studied fauna are of Late Mississippian (Chesterian; early

Serpukhovian) age from the Slade Formation. The Slade Formation includes several members (Figure 2.1), but the most fossiliferous and echinoderm-rich of these is the

Ramey Creek Member, from which both fossil faunas were collected. The Ramey Creek

Member conformably overlies the Tygarts Creek Member and is overlain conformably by the Maddox Branch Member (Ettensohn et al., 1984) (Figure 2.1). Figure 2.1 shows a generalized stratigraphic section for Upper Devonian, Mississippian and Lower

Pennsylvania rocks in northeastern Kentucky, as well as the stratigraphic position of the

Ramey Creek Member.

Figure 3.1 shows the stratigraphic section at the Valley Stone Quarry, Carter

County, one of the collection sites (Lierman et al., 2011), and Figure 3.2 shows a schematic drawing of the same section. The main lithology of the Ramey Creek Member is a thin-bedded, grayish-green, argillaceous calcarenite with interbedded shale

(Ettensohn et al., 1984), but local lenses of well-washed, bioclastic calcarenites are common. Chert nodules and red-chert-replaced fossils only occur in the Ramey Creek

Member (Figure 3.3). The Ramey Creek Member generally differs from the underlying limestones of Tygarts Creek by its thinly bedded nature and the presence of shale, chertified fossils, and fine-grained, argillaceous and limestones (Ettensohn et al., 1984)

(Figures 1.5 and 3.2).

Figure 3.1. Units exposed at the Valley Stone Quarry, Carter County, KY (from Lierman et al., 2011). Collections in this study were made from the Ramey Creek Member.

Figure 3.2. Schematic stratigraphic section from the Valley Stone Quarry, Carter County, Kentucky, study area. The Ramey Creek Member and its fossils are the focus of this study (from Lierman et al., 2011).

Figure 3.3. Specimen with chert replacement of echinoid specimens at the Valley Stone Quarry, Carter County, KY.

In the Valley Stone Quarry, four distinct Ramey Creek lithofacies were recognized (Figure 3.4): 1) a white, well-washed, thick-bedded, coarse-grained calcarenite with prominent crossbedding and occasional ripple marks (Figures 3.4, 3.5,

3.6, and 3.7); 2) a gray, coarse-grained, thin-bedded, argillaceous calcarenite with sparse interbedded grey-green shale partings (Figure 3.8); 3) a gray, fine-grained, thin-bedded, calcarenite and interbedded calcilutite (Figure 3.9); and 4) a gray, thin-bedded,

argillaceous calcilutite with thin gray-green shale partings (Figure 3.10). As already noted, these lithofacies apparently occur randomly across an area of approximately 0.17 km² (0.07 mi²) (Figure 1.1), and they all contain fossils that include echinoderms. In the coarse-grained calcarenite lithofacies, most of the fossils are imperfectly preserved along stylolitic bedding planes (Figure 3.11). In contrast, preservation is substantially better on the bedding planes of the finer-grained lithofacies (Figure 3.10). Although the various

Ramey Creek Member lithofacies are shown as superimposed subunits in Figure 3.4, according to Walther’s Law, at some time and place during Ramey Creek Member deposition, they would have been contemporaneous.

Figure 3.4. Ramey Creek Member lithofacies on U.S. 60 southwest of the Valley Stone Quarry (see Figure 1.6). The fine dashed lines in the coarse-grained calcarenite lithofacies outline major bedforms.

Figure 3.5. Example of low-angle crossbedding and planar bedding from the coarse- grained calcarenite lithofacies at the Valley Stone Quarry.

Figure 3.6. Sectional view of interference ripples (red, dashed line) from the fine-grained calcarenite and interbedded calcilutite lithofacies at the Valley Stone Quarry.

Figure 3.7. Sectional view of the coarse-grained calcarenite lithofacies from the Valley Stone Quarry with two Composita brachiopod shells.

Figure 3.8. Slab from the Valley Stone Quarry showing specimens on the coarse-grained, argillaceous calcarenite lithofacies.

Figure 3.9. Slab from the Valley Stone Quarry showing brachiopod, crinoid (black arrow), edrioasteroid and ophiuroid (black arrow) specimens on a hardground bedding- plane surface from the fine-grained calcarenite and interbedded calcilutite lithofacies. Pits in the surface of the specimen probably reflect corrosion during exposure on the bottom. The arms of the ophiuroid may reflect preferential current orientation.

Figure 3.10. Slab from the Valley Stone Quarry showing a buried crinoid (black arrow) specimen from the argillaceous calcilutite lithofacies.

Figure 3.11. Slab from the Valley Stone Quarry showing brachiopod and crinoid specimens (black arrorws) on a stylolitic bedding plane from the coarse-grained calcarenite lithofacies.

In the 213 Quarry (Figure 3.12 and 3.13), the entire fossil-bearing parts of one layer were collected. This layer is a very thin- to thin-bedded (0.7–7.5 cm) (Figure 3.14), argillaceous calcarenite with interbedded gray-green shales (Figures 1.7, 3.12, 3.13, and

3.14). It is similar to the calcarenite and interbedded calcilutite lithofacies at the Valley

Stone Quarry. The Ramey Creek Member has more limestones and is coarser than the

Maddox Branch Member. The Ramey Creek Member also has chert nodules and chert replacement of fossils (Figure 1.2 and 3.3).

Locally abundant fossils with echinoderms occur on both the bottoms and tops of the slabs (Figures 1.7, 3.3, 3.15). In a few places, a horizon of breccia and deformed limestone sits just below fossils on the bottom of the slabs (Figures 3.13, 3.16, 3.17).

Figure 3.18 shows a cross sectional view of the top-of-slab and bottom-of-slab community with examples of graded bedding, a gutter cast and cross-bedding.

Figure 3.12. Units exposed at the 213 Quarry, Powell County, Ky. Fossils for this study were collected from the very top of the Ramey Creek Member in this picture (see Figure 3.13).

Figure 3.13. Close-up view of the Ramey Creek Member at the 213 Quarry (see Figure 3.9), showing the single lithofacies and thin-bedded nature of the unit. The fossils from this quarry were collected just above the breccia unit at the top.

Figure 3.14. Photo of the thin-bedded nature of some slabs from the 213 Quarry. Fossil assemblages occur on the top and bottom of this slab. The broken face of the slab is outlined in red dashes.

Figure 3.15. Preservation and abundance of fossils on the top-of-slab, from the 213 Quarry. Note the crude rectangle orientations of Archimedes spires and crinoid (red dashed line).

Figure 3.16. Photo of breccia and deformed limestone at the base of a slab from the 213 Quarry.

Figure 3.17. Cross sectional view through a slab, showing breccia at the base.

Figure 3.18. Cross sectional view of the top-of-slab and bottom-of-slab communities, showing the graded bedding, a gutter cast and cross-bedding associated with a likely tempestite or storm layer.

3.2. History of Stratigraphic Nomenclature

According to Ettensohn et al. (1984), the names Newman Limestone and

Pennington Shale (Figure 3.19) were introduced into east-central Kentucky by Campbell

(1898a, b), who first formalized them in a study of the Big Stone Gap coal field in southwestern Virginia. The formations were described from exposures at Big Stone Gap on Pine Mountain.

The Newman Limestone was described as having an upper sandstone and shale unit, a middle limestone and shale unit, and a basal unit. However, Campbell (1898a, b) did not clearly define the Newman Limestone or the Pennington Formation in east-central

Kentucky, except to describe the Newman as a limestone and the Pennington as a shale unit containing thin beds of limestone. Ettensohn et al. (1984) solved this problem by redefining and distinguishing new units in the Cumberland escarpment outcrops

Butts’ (1922) work included the Mississippian stratigraphy in eastern and western

Kentucky. He described east-central Kentucky units and correlated them mostly with western Kentucky and southern Illinois units. The studies of Stokley (1949), Stokley and

McFarlan (1952), Stokley and Walker (1953), McFarlan and Walker (1956), and

Patterson and Hosterman (1961) were refinements in the division and correlation of

Butts’ original units.

Horne and others (1971, 1974) described the Mississippian rock section in northeastern Kentucky in terms of the Lee-Newman barrier-shoreline depositional model.

This highly controversial model was contested on lithostratigraphic and biostratigraphic grounds by Ettensohn and Dever (1975, 1979), Ettensohn (1975a, 1977, 1980, 1981a, b),

Ettensohn and Peppers (1979), Rice and others (1979), and Englund and Henry (1981) and was finally resolved through the redefinition and renaming of units (Ettensohn et al.,

1984).

Figure 3.19. The stratigraphic nomenclature of several authors in east-central Kentucky showing subdivisions of the Slade and Paragon formations (adapted from Ettensohn et al., 1984).

3.3. Slade Formation

The Slade Formation is named for the section of limestone, dolostone, and minor shale exposed in the southwestern highwall of the Natural Bridge Company quarry near the Mountain Parkway, about 6.5 km west-northwest of Slade, Kentucky (Ettensohn et al., 1984). The Slade Formation comprises twelve members and one bed. From oldest to youngest, these include the Renfro, St. Louis, Ste. Genevieve, Warix Run and Mill Knob members, the Cave Branch Bed, and the Armstrong Hill, Holly Fork, Rosslyn, Tygarts

Creek, Ramey Creek, Maddox Branch, and Poppin Rock members (Figures 2.1, and 3.2).

This study is focused on the Ramey Creek Member in the upper part of the Slade

Formation.

3.4. Upper Part of Slade Formation

The upper Slade stratigraphic succession consists of thin- to thick-bedded limestones with some shales and dolostones. The upper Slade consists of one bed and seven members: the Cave Branch bed, and the Armstrong Hill, Holly Fork, Rosslyn,

Tygarts Creek, Ramey Creek, Maddox Branch, and Poppin Rock Members (Figures 2.1,

3.1, and 3.2).

3.4.1. Poppin Rock Member

The Poppin Rock Member is mostly composed of thin- to thick-bedded, crystalline calcarenite. Dolostones occur locally in the southern parts of the outcrop belt.

The lower parts of the Poppin Rock Member represent deposition in a carbonate sandbelt environment, whereas upper parts represent deposition in a shallower-water, back-

sand-belt lagoonal setting (Ettensohn 1974, 1975b, 1977, 1980; Chesnut and Ettensohn,

1988).

3.4.2. Maddox Branch Member

The Maddox Branch Member is composed mainly of green calcareous shale. This member has very limited fossil diversity and was deposited in a relatively deep, outer- platform, open-marine environment (Ettensohn 1974, 1975b, 1977, 1980).

3.4.3. Ramey Creek Member

The Ramey Creek Member is mostly grayish-green, argillaceous calcarenite and interbedded shale. The member is highly burrowed, fossiliferous and it contains the most diverse fauna of any member in the upper part of the Slade Formation. Brachiopods, bryozoans, rugose corals, sponges, and echinoderms are the dominant fossils in this member. The Ramey Creek Member represents deposition in a shallow open-marine environment (Ettensohn 1974, 1975b, 1977, 1980).

3.4.4. Tygarts Creek Member

The Tygarts Creek Member is the most widespread member of the upper part of the Slade Formation. The member is largely composed of white, medium to coarsely crystalline, bioclastic to oolitic calcarenite. Channeling, wave ripples, and large intraclasts are prominent in this member. Fossil diversity is extremely low because of the high-energy setting. Fossils, which have been recorded, include large, thick-shelled gastropods and stemless crinoids predominate (Ettensohn, 1975b). This member

represents deposition in an agitated, carbonate sand-belt environment (Ettensohn 1974,

1975b, 1977, 1980).

3.4.5. Rosslyn Member

The Rosslyn Member is thin and largely calcarenite. This member is bioturbated and contains dolomitic mud chips, shale fragments, rounded clasts, fossil fragments. It represents deposition on back-sand-belt intertidal flats and contains a high density of trace fossils (Ettensohn, 1975b).

3.4.6. Holly Fork Member

The Holly Fork Member is an upward-fining, dolomitic, tidal-flat and tidal- channel sequence. Vertical burrows, channels, birdseyes and mud cracks characterize the member; fossils are rare. The member represents deposition on carbonate tidal flats

(Ettensohn, 1975b, 1977, 1980).

3.4.7. Armstrong Hill Member

The Armstrong Hill Member consists of thin- to thick-bedded, gray calcilutite with thin shale partings. The uppermost parts are dolomitic. This member contains abundant burrows and a predominantly molluscan fauna of gastropods and pelecypods. It represents an open, channel-lagoon deposit with locally restricted areas (Ettensohn, 1974,

1975b, 1977).

3.4.8. Cave Branch Bed

The only formally recognized bed in the upper Slade Formation consists of red and green silty shale and mudstone that is interbedded with thin calcilutite lenses. Except

for algae, fossils are very rare (Ettensohn et al., 1984). The bed represents terrigenous terra rossa accumulation on underlying carbonates and its reworking as red muds on intertidal mudflats. Mud cracks are the most common sedimentary structure.

3.5. Depositional environments

The upper Slade, which includes the studied Ramey Creek Member, represents a transgression followed by a regression (Figure 2.1 and 3.1). The Ramey Creek Member represents a shallow, open-marine facies in the transgressive parts of the upper Slade

Formation. The member is interpreted in the context of the Shaw-Irwin model, using

Walther’s Law (Figures 3.20 and 3.21).

The Shaw (1964) and Irwin (1965) models are commonly used to interpret clear- water, epeiric-shelf sedimentation specific to shallow-water environments in very gently dipping cratonic sea floor (Figure 3.20). Figure 3.21 places upper Slade members in their likely depositional continuum from the Shaw-Irwin models. The Ramey Creek and

Maddox Branch members represent open-marine deposition seaward of the Tygarts Crek sand belt (Figure 3.21). These members are interpreted to have been deposited below normal wave base (~10 meters) but above typical storm wave base (~70 meters) because of stratigraphic position and lithologic and fossil content.

Figure 3.20. Distribution of environments in the generalized Shaw (1964) and Irwin (1965) models.

The study area is interpreted to have been in a Paleozoic storm belt based on paleogeographic reconstructions (Marsaglia and Klein, 1983; Duke, 1985). Storms would have agitated and reworked sediment from higher and more shoreward environments, and sent clouds of fine muddy debris seaward to settle out in parts of shallow open-marine or deeper-water settings (Aigner, 1982). Coarser debris were reworked in place or transported seaward by storm-related backflow currents (Aigner, 1982; Kreisa and

Bambach, 1982), which generated local shoal-like deposits interspersed with muddy basinal lows between (Ettensohn et al., 1984, 2004).

Figure 3.21. The stratigraphic section at the Valley Stone Quarry shown as parts of a vertical stratigraphic sequence (A) and as part of a lateral environmental, Shaw-Irwin succession (B). Walther’s Law explains how a lateral succession of environments becomes a vertical stratigraphic sequence. The studied Ramey Creek Member represents deposition in a shallow open-marine environment (red B=local basins; red S=local shoals) (Ettensohn et al., 1984).

CHAPTER 4: FOSSIL-ASSEMBLAGE TAXONOMY

In order to interpret the relationships among the organisms of a community, it is necessary to firmly understand the taxonomic makeup of the assemblages being studied

(Whittington, 1964; Fagerstrom, 1964). Although taxonomy is not the sole basis on which paleoecological interpretations rest, its importance stems from the fact that paleoecology is about fossils, and it is important to know the specific identity of the fossils that are present and what each species was like in life. Hence, the numbers of individual taxa on the rock slabs from the 213 Quarry and Valley Stone Quarry localities were recorded, as were all loose specimens from both sections. In addition, characteristics of each species or genus, along with the number of individuals, were noted, tallied and presented in Tables 4.1–4.14. These tallies will be the basis for statistical analyses and comparison of the communities in Chapter 5. With a few exceptions, bryozoans were only identified to the genus level based on general characteristics, as species-level identification would require much detailed thin-section work, which was beyond the scope of this study. In addition, with the exception of

Archimedes, where each spire was tallied as an individual colony, each bryozoan fragment was tallied as an individual colony, meaning that bryozoan “individuals,” other than Archimedes, are probably overrepresented in the tallies. Similarly, because brachiopods are generally represented by two valves, any two valves were considered to be “individuals,” except in those cases where the valves were articulated.

This study was undertaken in part because echinoderms, which are normally rare in the fossil record, are relatively common in the two faunas studied, and hence, provide

an opportunity to understand and compare the communities and their taphonomy.

Crinoids are the dominant echinoderm class in these communities. What makes these faunas so unusual is the fact that most of the crinoid calyces, which are necessary for identification, are complete and fully articulated. Several intact thecae from other echinoderm groups are also well preserved. This kind of preservation provides an unusual opportunity to examine the echinoderms individually as species and as parts of ecological communities.

The following sections describe a new encrinasterid ophiuroid species, previously known species and genera, and census-population data. Specimen tallies from each locality and community are included and discussed in remaining parts of this chapter.

4.1. Systematic paleontology of the new ophiuroid species

In general, preservation was sufficient to recognize most of the ophiuroid species as having already been described. In a few cases, however, the silicification prevented the identification of species, and fossils could only be identified to class, ordinal, or genus level. One exception is a new ophiuroid species in the genus Schoenaster, which is described below. This species will be formally described in a separate publication.

Class Ophiuroidea Gray, 1840

Order Oegophiurida Matsumoto, 1915

Suborder Lysophiurina Gregory, 1896

Family Encrinasteridae Schuchert, 1914

Genus Schoenaster Meek and Worthen, 1866

Palasterina (Schoenaster) Meek and Worthen, 1860, p. 449.

Schoenaster Meek and Worthen, 1866, p. 277; Schuchert, 1915, p. 202.

Encrinaster Spencer, 1930, p. 418 (partim).

? Euzonosoma Spencer and Wright, 1966, p. U86 (partim).

Type species.–Palasterina (Schoenaster) fimbrata Meek and Worthen 1860, from the Mississippian St. Louis Limestone of St. Clair County, Illinois.

Diagnosis.–Small to large, pentagonal, asteroid-like individuals with arms that taper uniformly to acute points; arms never petaloid. Arms and dorsal surface are convex, covered with small, tumid, polygonal plates with granular ornament that extend to arm tips. Disc is large with well-developed v-shaped interrays containing small, polygonal, imbricating plates. Margins of disc between rays is concave, with or without differentiated ambital framework (marginal) plates. Ventral plates may show projecting spines, but their absence may be a preservational effect. Ambulacrals (Ambb) alternate and are L- or boot-shaped with an elongate “toe” parallel to arm axis. Laterals (LL) subventral with broad oral faces, rectangular with an adradial termination near Ambb

(Figures 4.1–4.2; 4.6–4.7); they are arranged with their long axes directed obliquely outward, giving a twisted-rope appearance, after which the genus was named. Mouth frame robust, formed from pairs of proximal Ambb in each ray that join interradially with adjacent pairs of proximal Ambb to form a star-shaped frame around the mouth.

Remarks.–As originally designated, Meek and Worthen (1860) placed their specimens in the asteroid genus Palasterina M’Coy, 1851, but the LL in their specimens were oriented obliquely to the Ambb, which differs from the perpendicular orientation to the Ambb in Palasterina. Hence, they created the subgenus Schoenaster to accommodate

this difference. However, by 1866, Meek and Worthen thought that this difference and a few others were substantial enough to separate specimens with the perpendicular orientation as a distinct genus under the name Schoenaster. Subsequently, some species of the genus were synonymized with Encrinaster (Spencer, 1930) and Euzonosoma

(Spencer and Wright, 1966) based on the presence of ventral-surface spines and arm shape, respectively. Jell (1997) resurrected the genus based on the absence of petaloid arms. More on the history of the genus Schoenaster is presented by Harper and Morris

(1978) and Jell (1997).

The combination of long, distally tapering arms and concave disc margins readily differentiate Schoenaster from other encrinasterid ophiuroids. Ventral-surface spines may also be a distinguishing trait, but their presence or absence apparently depends on preservational state.

Schoenaster n. sp.

Figures 4.1–4.7

Diagnosis.–Species characterized by thick, stout, L-shaped Ambb in proximal parts of oral arm surfaces becoming rectangular distally (Figure 4.1). Concave ambital margins with poorly to moderately developed framework plates (Figure 4.2). Ventral-surface spines apparently absent. Aboral surface composed of many irregular, pustulose plates with a raised, carinal ridge along the mid-line of each ray that terminates in an irregular boss (Figures 4.3 and 4.4) above the mouth frame on the aboral disc.

Description.–Small to large, pentagonal individuals with a cross-section in life that was low and shield-like. Dorsal surface covered with small, pustulose, polygonal plates

(Figures 4.3 and 4.4). Arms taper uniformly to acute points (Figure 4.5); length of arms

beyond disc range from 0.2–6.0 cm, averaging 1.7 cm. In well-preserved specimens, a carinal ridge, one-to-two-plates wide, extends along the mid-line of each ray and terminates in a prominent boss (Figure 4.4) above the mouth frame on the dorsal surface.

Disc is pentagonal and concave at ambitus (Figures 4.2, 4.4 and 4.5) with a perimeter that ranges from 0.5–6.5 cm, averaging 2.6 cm; sides range from 0.1–1.3 cm in length, averaging 0.6 cm. Ambital framework plates poorly to moderately differentiated (Figure

4.2). Ambb and LL ossicles stout and robust (Figures 4.1 and 4.6–4.7). Ambb form a biserial row with plates on either side of the perradial suture alternating; perradial suture straight or gently undulatory (Figures 4.1 and 4.6). Ambb are “L”- or boot-shaped with elongate part parallel to arm axis and are concave on distal and abradial margins (Figure

4.6); Ambb become rectangular distally and more blocky proximally toward mouth frame

(Figures 4.1 and 4.6). As per the genus, Laterals (LL) subventral and robust with broad oral faces; they are rectangular with pointed adradial terminations near Ambb and arranged with their long axes directed obliquely outward (Figures 4.1 and 4.6). The “L”- shape of the Ambb may be difficult to observe, because adjacent LL articulate with

Ambb at the internal angle of the “L” (Figures 4.1 and 4.6). LL may have functioned to close and protect the ambulacral groove (Figure 4.7). Podial gap shared equally by adjacent Ambb and LL (Figure 4.1). Mouth frame formed from proximal pairs of adjacent elongate Ambb in each ray (=circumorals?) that join interradially (Figures 4.2,

4.5 and 4.6). Spines and pustules on plates apparently absent from ventral surface.

Remarks.–This new species differs from the Meramecian (St. Louis) type, S. fimbratus, which has very small, delicate Ambb and less elongate, squat LL, in its stout, thicker, almost rectangular, Ambb and more elongate LL. The new species also apparently lacks

the ventral spines that characterize S. fimbratus. The new species differs from the

Osagean (Burlington) species, S. wachsmuthi, in that S. wachsmuthi has more ovoid and less oblique LL. Finally, the new species differs from the Kinderhookian species, S. legrandensis, in that S. legrandensis has nearly straight, narrow arms and oblong LL that are rounded at the ends. The genus Schoenaster was previously known only from Lower and Middle Mississippian (Kinderhookian–Meramecian) rocks, but the new occurrence described herein extends its range into Late Mississippian (Chesterian) time.

Figure 4.1. Camera lucida drawing of the arm (oral surface) of Schoenaster n. sp., showing the thick, stout nature of the ambulacrals (Ambb), the laterals (LL), podial gaps, and perradial suture. L-shaped nature of Ambb cannot be seen because LL are articulating with Ambb.

Figure 4.2. Camera lucida drawing of the oral surface of Schoenaster n. sp., showing the poorly differentiated nature of the ambital framework and the nature of the oral framework plates that derive from the proximal Ambb. Coarse stipple, Ambb; fine stipple, oral framework plates; random stipple, rock matrix.

Figure 4.3. Dorsal surface of an arm, showing small, pustulose, polygonal plates that comprise the dorsal surface (magnification X27).

Figure 4.4. Dorsal view of Schoenaster n. sp. showing the pustulose dorsal surface, carinal ridges, and dorsal bosses.

Figure 4.5. A typical ventral surface of Schoenaster n. sp., showing tapering arms and concave ambital margins. Oral framework plates are visible in center of disc (magnification X10). 59

Figure 4.6. Ventral surface of Schoenaster n. sp. photographed under water, showing prominent lateral and ambulacral ossicles. Note the stout, robust nature of the L-shaped Ambb (one Amb is outlined in red), the robust nature of the LL, and the poorly developed nature of the ambital framework plates (magnification X15).

Figure 4.7. Distal arm segment of Schoenaster n. sp. under water, showing closure by the lateral plates to protect the ambulacral groove (magnification X20). 60

4.2. Quarry Taxa

Before statistical metrics can be applied for comparative purposes, a complete list of taxa, their numbers, their presence or absence at the two localities, and their included lithofacies must be compiled. That basic information is provided below in Tables 4.1–4.9 by locality and lithofacies.

4.2.1. The 213 Quarry

At the 213 Quarry study area in Powell County, Kentucky, a total of forty-three invertebrate genera and two vertebrate form genera were identified, and many of these were identifiable to the species level. The most abundant in terms of species numbers are the crinoids followed by the brachiopods and bryozoans. However, in terms of absolute numbers of individuals, bryozoan colonies far outnumber individual species in any other taxon. Sixteen species of crinoids, eight species of brachiopods and nine genera/species of bryozoans are present. Species of lesser abundance include two echinoid species, one polychaete worm tube, one blastoid, one pelecypod, two cnidarians and various vertebrate form genera. Table 4.1 shows the complete list of genera/species for the 213

Quarry study area. From the 213 Quarry specimens, eight phyla, one of which includes the Chordata (Subphylum Vertebrata), twelve classes, and forty-five species represented.

Slabs from the 213 Quarry were further analyzed by examining specimens on the tops and bottoms of the slabs.

Table 4.1 Complete list of genera/species for the 213 Quarry study area.

Constituent Genera/Species for the 213 Phylum Class Quarry Anthozoa Zaphrentoides spinulosus Cnidaria Scyphozoa? Sphenothallus sp. Mollusca Pelecypoda Sulcatopinna missouriensis Annelida? Polychaeta? Crinicaminus haneyensis Arthropoda Ostracoda Amphissites?, Bairdia?, Coronakirkbya? Anthracospirifer leidyi, Cleiothyridina sublamellosa, Composita lewisensis, Articulata Composita subquadrata, Diaphragmus Brachiopoda elegans, Eumetria verneuliana Inarticulata Oehlertella pleurites, Orbiculoidea sp. Archimedes cf. meekanus, Eridopora sp., Fenestella sp., Glyptopora punctipora, Bryozoa Stenolaemata Lyroporella sp., Polypora sp., Rhombopora sp., Septopora sp., Sulcoretepora sp. Blastoidea Pentremites elegans Lepidesthes formosa, Archaeocidaris Echinoidea megastylus (isolated spines only) Acrocrinus shumardi, Agassizocrinus lobatus (juvenile), Anartiocrinus maxvillensis, Aphelecrinus mundus, Echinodermata Camptocrinus cirrifer, Cryphiocrinus girtyi, Cymbiocrinus grandis, Crinoidea Cymbiocrinus tumidus, Dasciocrinus florealis, Pentaramicrinus gracilis, Phanocrinus bimagnaramus, Phanocrinus maniformis, Pterotocrinus acutus, Taxocrinus whitfieldi, Tholocrinus spinosus, Zeacrinites magnoliaeformis Chondrichthyes Chordata Cladodus, Venustodus (teeth)

4.2.1.1. The 213 Quarry: Top of Slabs

The 213 Quarry records two different communities, one preserved on top of the slab and an earlier community now preserved on the bottom of the slab. The taxa on the top of the slab are noted in Table 4.2. Any loose samples were included in specimen counts for the top. An “X” denotes species that were present, and the “A” denotes species that were absent.

Table 4.2 Complete list of genera/species from the tops of the slabs in the 213 Quarry study area.

213 Quarry Census Top of Summary Slabs Anthozoa (1) Zaphrentoides spinulosus X

Scyphozoa? (1) Sphenothallus sp. X

Pelecypoda (1) Sulcatopinna missouriensis X

Polychaeta? (1) Crinicaminus haneyensis X

Ostracoda (3) Amphissites? X Bairdia? X Coronakirkbya? X

Articulata (6) Anthracospirifer leidyi X Cleiothyridina sublamellosa X Composita lewisensis X Composita subquadrata X

Table 4.2 (Continued)

Diaphragmus elegans X Eumetria verneuliana X

Inarticulata (2) Oehlertella pleurites X Orbiculoidea sp. X

Stenolaemata (9) Archimedes cf. meekanus X Eridopora sp. X Fenestella sp. X Glyptopora punctipora X Lyroporella sp. X Polypora sp. X Rhombopora sp. X Septopora sp. X Sulcoretepora sp. X

Blastoidea (1) Pentremites elegans X

Echinoidea (2) Lepidesthes formosa X Archaeocidaris megastylus A (isolated spines only)

Crinoidea (16) Acrocrinus shumardi X Agassizocrinus lobatus X (juvenile) Anartiocrinus maxvillensis X Aphelecrinus mundus X Camptocrinus cirrifer X Cryphiocrinus girtyi X Cymbiocrinus grandis X Cymbiocrinus tumidus X Dasciocrinus florealis X Pentaramicrinus gracilis X

Table 4.2 (Continued)

Phanocrinus bimagnaramus X Phanocrinus maniformis X Pterotocrinus acutus X Taxocrinus whitfieldi X Tholocrinus spinosus X Zeacrinites magnoliaeformis X

Chondrichthyes (2) Cladodus X Venustodus X

Species Present/Common 44 (X): Species Absent (A): 1

4.2.1.2. The 213 Quarry: Bottom of Slabs

The taxa from the bottoms of the slabs are listed in Table 4.3. An “X” denotes species that were present, and an “A” denotes species that were absent.

Table 4.3 Complete list of genera/species from the bottoms of the slabs in the 213 Quarry study area.

213 Quarry Census Bottom Summary of Slabs Anthozoa (1) Zaphrentoides spinulosus X

Scyphozoa? (1) Sphenothallus sp. X

Pelecypoda (1) Sulcatopinna missouriensis A

Table 4.3 (Continued)

Polychaeta? (1) Crinicaminus haneyensis A

Ostracoda (3) Amphissites? A Bairdia? A Coronakirkbya? A

Articulata (6) Anthracospirifer leidyi X Cleiothyridina sublamellosa X Composita lewisensis X Composita subquadrata X Diaphragmus elegans X Eumetria verneuliana A

Inarticulata (2) Oehlertella pleurites A Orbiculoidea sp. X

Stenolaemata (9) Archimedes cf. meekanus X Eridopora sp. X Fenestella sp. X Glyptopora punctipora X Lyroporella sp. A Polypora sp. X Rhombopora sp. X Septopora sp. X Sulcoretepora sp. X

Blastoidea (1) Pentremites elegans X

Echinoidea (2) Lepidesthes formosa A Archaeocidaris megastylus X (isolated spines only)

Table 4.3 (Continued)

Crinoidea (16) Acrocrinus shumardi A Agassizocrinus lobatus A (juvenile) Anartiocrinus maxvillensis A Aphelecrinus mundus A Camptocrinus cirrifer X Cryphiocrinus girtyi A Cymbiocrinus grandis X Cymbiocrinus tumidus A Dasciocrinus florealis X Pentaramicrinus gracilis X Phanocrinus bimagnaramus X Phanocrinus maniformis A Pterotocrinus acutus X Taxocrinus whitfieldi X Tholocrinus spinosus A Zeacrinites magnoliaeformis A

Chondrichthyes (2) Cladodus A Venustodus A

Species Present/Common 25 (X): Species Absent (A): 20

4.2.2. Valley Stone Quarry

The Valley Stone Quarry in Carter County, Kentucky, had a larger diverse species count than the 213 Quarry. Sixty-five genera were identified, and many of these were identifiable to the species level. The most abundant taxa were crinoids, followed by brachiopods and bryozoans. Eighteen species of crinoids, thirteen species of brachiopods and ten genera/species of bryozoans are present in the fauna. In addition, two species of edrioasteroids, one species of blastoid, one echinoid, one ophiuroid, one pelecypod, one

scyphozoan, one sponge, one tabulate coral, and one possible scyphozoan cnidarian are present in the fauna. Table 4.4 shows the complete list genera/species for Valley Stone

Quarry study area. From the Valley Stone Quarry, seven invertebrate phyla and one chordate subphylum, thirteen classes, and sixty-five species are present. Slabs and specimens from the Valley Stone Quarry were categorized further by lithofacies. Rather than breaking the lithofacies down by tops or bottoms of slabs, the tops and bottoms of slabs from multiple beds were combined, and species were noted across four contiguous lithofacies: coarse-grained calcarenite, coarse-grained argillaceous calcarenite, fine- grained calcarenite and interbedded calcilutite, and argillaceous calcilutite. The four lithofacies were classified based on the Grabau (1903) classification system (Table 4.5).

Table 4.4 Complete list genera/species for Valley Stone Quarry by phylum and class.

Constituent Genera/Species for Phylum Class Valley Stone Quarry Porifera Demospongea Belemnospongia fascicularis Anthozoa Unidentifiable tabulate coral? Cnidaria Sphenothallus sp., Scyphozoa? Paraconularia missouriensis Mollusca Pelecypoda Sulcatopinna missouriensis

Annelida? Polychaeta? Crinicaminus haneyensis Anthracospirifer leidyi, Cleiothyridina sublamellosa, Composita lewisensis, Composita subquadrata, Diaphragmus elegans, Articulata Dielasma sp., Eumetria costata, Brachiopoda Eumetria verneuliana, Orthotetes kaskaskiensis, Punctospirifer sp., Streptorhynchus sp. Inarticulata Lingulipora sp., Orbiculoidea sp.

Table 4.4 (Continued) Anisotrypa sp., Archimedes cf. meekanus, Eridopora sp., Fenestella sp., Fenestrellina sp., Polypora sp., Bryozoa Stenolaemata Rhombopora sp., Septopora sp., Thamniscus? Unidentifiable encrusting bryozoan Unidentifiable edrioasteroid, Edrioasteroidea Lepidodiscus laudoni Blastoidea Pentremities elegans Echinoidea Archaeocidaris megastylus Ophiuroidea Unidentifiable encrinasterid Agassizocrinus lobatus, Anartiocrinus maxvillensis, Aphelecrinus mundus, Camptocrinus cirrifer, Cryphiocrinus girtyi, Echinodermata Cymbiocrinus grandis, Cymbiocrinus tumidus, Dasciocrinus florealis, Eupachycrinus boydii, Crinoidea Linocrinus wachsmuthi, Pentaramicrinus gracilis, Phacelocrinus longidactylus, Phanocrinus bimagnaramus, Phanocrinus maniformis, Platycrinites sp., Talarocrinus sp., Taxocrinus whitfieldi, Tholocrinus spinosus Agassizodus, Cladodus, Ctenacanthus, Deltodus, Chondrichthyes Deltodopsis, Listracanthus?, (Teeth) Lophodus, Mesodmodus, Chordata Psammodus?, Polyrhizodus, Taeniodus?, Venustodus Chondrichthyes (dermal skin Petrodus plates)

Table 4.5 The four lithofacies at the Valley Stone Quarry based on the Grabau (1903) classification.

Grabau (1903) Classification Rock Type Description (Grain size) Calcirudite More than 50%, grains coarser than sand Calcarenite More than 50%, sand-sized grains Calcisiltite More than 50%, silt-sized grains Calcilutite More than 50%, clay-sized grains

4.2.2.1. Valley Stone Quarry: Coarse-grained calcarenite

The coarse-grained calcarenite lithofacies consists of well-washed, bioclastic, sand-size grains. An “X” denotes species that were present and the “A” denotes species that were absent in the coarse-grained calcarenite lithofacies (Table 4.6).

Table 4.6 Complete combined list of genera/species for Valley Stone Quarry coarse- grained calcarenite lithofacies by class.

Coarse- Valley Stone Quarry Census grained, Summary calcarenite Demospongea (1) Belemnospongia fascicularis X

Anthozoa (1) Unidentifiable tabulate coral? X

Scyphozoa? (2) Sphenothallus sp. X

Table 4.6 (Continued)

Paraconularia missouriensis X

Pelecypoda (1) Sulcatopinna missouriensis X

Polychaeta? (1) Crinicaminus haneyensis X

Articulata (11) Anthracospirifer leidyi X Cleiothyridina sublamellosa A Composita lewisensis X Composita subquadrata X Diaphragmus elegans X Dielasma sp. A Eumetria costata X Eumetria verneuliana X Orthotetes kaskaskiensis X Punctospirifer sp. A Streptorhynchus sp. X

Inarticulata (2) Lingulipora sp. X Orbiculoidea sp. X

Stenolaemata (10) Anisotrypa sp. X Archimedes cf. meekanus X Eridopora sp. A Fenestella sp. A Fenestrellina sp. A Polypora sp. X Rhombopora sp. X Septopora sp. A Thamniscus? X Unidentifiable encrusting bryozoan X

Edrioasteroidea (2)

Table 4.6 (Continued)

Unidentifiable edrioasteroid X Lepidodiscus laudoni A

Blastoidea (1) Pentremites elegans X

Echinoidea (1) Archaeocidaris megastylus X

Ophiuroidea (1) Unidentifiable encrinasterid brittle X star

Crinoidea (18) Agassizocrinus lobatus X Anartiocrinus maxvillensis X Aphelecrinus mundus X Camptocrinus cirrifer A Cryphiocrinus girtyi A Cymbiocrinus grandis X Cymbiocrinus tumidus X Dasciocrinus florealis X Eupachycrinus boydii A Linocrinus wachsmuthi X Pentaramicrinus gracilis X Phanocrinus bimagnaramus X Phanocrinus maniformis X Phacelocrinus longidactylus A Platycrinites sp. A Talarocrinus sp. X Taxocrinus whitfieldi X Tholocrinus spinosus X

Chondrichthyes (13) Agassizodus A Cladodus A Ctenacanthus A Deltodopsis A

Table 4.6 (Continued)

Deltodus A Listracanthus? A Lophodus A Mesodmodus A Petrodus A Psammodus? A Polyrhizodus A Taeniodus? A Venustodus A

Species Present/Common (X): 39 Species Absent (A): 26

4.2.2.2. Valley Stone Quarry: Coarse-grained, argillaceous calcarenite

The coarse-grained, argillaceous calcarenite lithofacies consists of sand grains that also includes clay. The “X” denotes species that were present and the “A” denotes species that were absent in the coarse-grained, argillaceous calcarenite lithofacies (Table

4.7).

Table 4.7. Complete combined list of genera/species for Valley Stone Quarry coarse- grained, argillaceous calcarenite lithofacies by class.

Coarse- grained, Valley Stone Quarry Census argillaceous Summary calcarenite Demospongea (1) Belemnospongia fascicularis X

Anthozoa (1)

Table 4.7 (Continued)

Unidentifiable tabulate coral? A

Scyphozoa? (2) Sphenothallus sp. X Paraconularia missouriensis A

Pelecypoda (1) Sulcatopinna missouriensis A

Polychaeta? (1) Crinicaminus haneyensis A

Articulata (11) Anthracospirifer leidyi X Cleiothyridina sublamellosa X Composita lewisensis X Composita subquadrata X Diaphragmus elegans X Dielasma sp. A Eumetria costata A Eumetria verneuliana X Orthotetes kaskaskiensis A Punctospirifer sp. X Streptorhynchus sp. A

Inarticulata (2) Lingulipora sp. A Orbiculoidea sp. X

Stenolaemata (10) Anisotrypa sp. A Archimedes cf. meekanus X Eridopora sp. A Fenestella sp. A Fenestrellina sp. A Polypora sp. X Rhombopora sp. X Septopora sp. A

Table 4.7 (Continued)

Thamniscus? X Unidentifiable encrusting bryozoan X

Edrioasteroidea (2) Unidentifiable edrioasteroid A Lepidodiscus laudoni A

Blastoidea (1) Pentremites elegans X

Echinoidea (1) Archaeocidaris megastylus A

Ophiuroidea (1) Unidentifiable encrinasterid brittle X star

Crinoidea (18) Agassizocrinus lobatus X Anartiocrinus maxvillensis A Aphelecrinus mundus X Camptocrinus cirrifer X Cryphiocrinus girtyi A Cymbiocrinus grandis X Cymbiocrinus tumidus X Dasciocrinus florealis X Eupachycrinus boydii A Linocrinus wachsmuthi X Pentaramicrinus gracilis A Phanocrinus bimagnaramus X Phanocrinus maniformis A Phacelocrinus longidactylus X Platycrinites sp. A Talarocrinus sp. A Taxocrinus whitfieldi A Tholocrinus spinosus A

Chondrichthyes (13)

Table 4.7 (Continued)

Agassizodus X Cladodus X Ctenacanthus A Deltodopsis A Deltodus A Listracanthus? A Lophodus A Mesodmodus A Petrodus A Psammodus? A Polyrhizodus A Taeniodus? A Venustodus X

Species Present/Common (X): 29 Species Absent (A): 36

4.2.2.3. Valley Stone Quarry: Fine-grained calcarenites and interbedded calcilutite The fine-grained calcarenite lithofacies consists of well-washed, bioclastic, fine sand-size grains. An “X” denotes species that were present and an “A” denotes species that were absent in the fine-grained calcarenite and interbedded calcilutite lithofacies

(Table 4.8).

Table 4.8 Complete combined list of genera/species for Valley Stone Quarry fine-grained calcarenites and interbedded calcilutite lithofacies by class.

Fine-grained calcarenite and Valley Stone Quarry Census interbedded Summary calcilutite Demospongea (1)

Table 4.8 (Continued)

Belemnospongia fascicularis X

Anthozoa (1) Unidentifiable tabulate coral? A

Scyphozoa? (2) Sphenothallus sp. X Paraconularia missouriensis A

Pelecypoda (1) Sulcatopinna missouriensis A

Polychaeta? (1) Crinicaminus haneyensis A

Articulata (11) Anthracospirifer leidyi X Cleiothyridina sublamellosa X Composita lewisensis A Composita subquadrata X Diaphragmus elegans X Dielasma sp. A Eumetria costata A Eumetria verneuliana A Orthotetes kaskaskiensis A Punctospirifer sp. A Streptorhynchus sp. A

Inarticulata (2) Lingulipora sp. A Orbiculoidea sp. X

Stenolaemata (10)

Table 4.8 (Continued)

Anisotrypa sp. A Archimedes cf. meekanus A Eridopora sp. A Fenestella sp. X Fenestrellina sp. X Polypora sp. X Rhombopora sp. A Septopora sp. A Thamniscus? X Unidentifiable encrusting bryozoan A

Edrioasteroidea (2) Unidentifiable edrioasteroid A Lepidodiscus laudoni A

Blastoidea (1) Pentremites elegans X

Echinoidea (1) Archaeocidaris megastylus X

Ophiuroidea (1) Unidentifiable encrinasterid brittle X star

Crinoidea (18) Agassizocrinus lobatus X Anartiocrinus maxvillensis X Aphelecrinus mundus A Camptocrinus cirrifer X Cryphiocrinus girtyi A Cymbiocrinus grandis A Cymbiocrinus tumidus A Dasciocrinus florealis X Eupachycrinus boydii A Linocrinus wachsmuthi X Pentaramicrinus gracilis A Phanocrinus bimagnaramus X

Table 4.8 (Continued)

Phanocrinus maniformis A Phacelocrinus longidactylus A Platycrinites sp. A Talarocrinus sp. X Taxocrinus whitfieldi A Tholocrinus spinosus A

Chondrichthyes (13) Agassizodus A Cladodus A Ctenacanthus A Deltodopsis A Deltodus A Listracanthus? A Lophodus A Mesodmodus A Petrodus A Psammodus? A Polyrhizodus A Taeniodus? A Venustodus A

Species Present/Common (X): 21 Species Absent (A): 44

4.2.2.4. Valley Stone Quarry: Argillaceous calcilutite

The argillaceous calcarenite lithofacies consists of a very fine-grained clay material. An “X” denotes species that were present, and an “A” denotes species that were absent in the argillaceous calcilutite lithofacies (Table 4.9).

Table 4.9 Complete combined list of genera/species for the Valley Stone Quarry argillaceous calcilutite lithofacies by class.

Valley Stone Quarry Census Argillaceous Summary calcilutites Demospongea (1) Belemnospongia fascicularis X

Anthozoa (1) Unidentifiable tabulate coral? A

Scyphozoa? (2) Sphenothallus sp. X Paraconularia missouriensis X

Pelecypoda (1) Sulcatopinna missouriensis X

Polychaeta? (1) Crinicaminus haneyensis A

Articulata (11) Anthracospirifer leidyi X Cleiothyridina sublamellosa X Composita lewisensis X Composita subquadrata X Diaphragmus elegans X Dielasma sp. X Eumetria costata X Eumetria verneuliana X Orthotetes kaskaskiensis X Punctospirifer sp. X Streptorhynchus sp. A

Inarticulata (2) Lingulipora sp. A Orbiculoidea sp. A

Table 4.9 (Continued)

Stenolaemata (10) Anisotrypa sp. X Archimedes cf. meekanus A Eridopora sp. X Fenestella sp. X Fenestrellina sp. A Polypora sp. X Rhombopora sp. X Septopora sp. X Thamniscus? X Unidentifiable encrusting bryozoan A

Edrioasteroidea (2) Unidentifiable edrioasteroid A Lepidodiscus laudoni X

Blastoidea (1) Pentremites elegans X

Echinoidea (1) Archaeocidaris megastylus X

Ophiuroidea (1) Unidentifiable encrinasterid brittle X star

Crinoidea (18) Agassizocrinus lobatus X Anartiocrinus maxvillensis X Aphelecrinus mundus X Camptocrinus cirrifer A Cryphiocrinus girtyi X Cymbiocrinus grandis X Cymbiocrinus tumidus X Dasciocrinus florealis X Eupachycrinus boydii X Linocrinus wachsmuthi X Pentaramicrinus gracilis X

Table 4.9 (Continued)

Phanocrinus bimagnaramus X Phanocrinus maniformis A Phacelocrinus longidactylus A Platycrinites sp. X Talarocrinus sp. A Taxocrinus whitfieldi X Tholocrinus spinosus A

Chondrichthyes (13) Agassizodus A Cladodus X Ctenacanthus X Deltodopsis X Deltodus X Listracanthus? X Lophodus X Mesodmodus X Petrodus X Psammodus? X Polyrhizodus X Taeniodus? X Venustodus A

Species Present/Common (X): 49 Species Absent (A): 16

4.3. Quarry Comparisons

The two study areas occur in the same stratigraphic unit and are approximately of the same age. Yet, they are similar and different in a variety of ways. Table 4.10 shows the occurrence of each species at each study area. An “X” means that the species is present for that quarry locality. The Valley Stone Quarry locality contained several species of brachiopods that were not observed at the 213 Quarry, such as, Dielasma sp.,

Lingulipora sp., Orthotetes kaskaskiensis, Punctospirifer sp., and Streptorhynchus sp. In

addition, various crinoid species are more abundant at the Valley Stone Quarry. These crinoids are present at the Valley Stone Quarry but not at the 213 Quarry: Cryphiocrinus girtyi, Cymbiocrinus tumidus, Linocrinus wachsmuthi, Phacelocrinus longidactylus,

Platycrinites sp., and Talarocrinus sp. Table 4.9, compares species occurrence at each quarry locality.

Table 4.10 Comparison of fauna from the 213 Quarry and Valley Stone Quarry by phylum and class. The “X’s” represent species that are present within each quarry.

213 Quarry vs. Valley Stone Quarry Taxa Valley Constituent 213 Stone Phylum Class Genera/Species Quarry Quarry Porifera Demospongea Belemnospongia fascicularis X Unidentifiable tabulate Anthozoa coral? X Cnidaria Zaphrentoides spinulosus X Sphenothallus sp. X X Scyphozoa? Paraconularia missouriensis X Mollusca Pelecypoda Sulcatopinna missouriensis X X Annelida? Polychaeta? Crinicaminus haneyensis X X Amphissites? X Arthropoda Ostracoda Bairdia? X Coronakirkbya? X Anthracospirifer leidyi X X Cleiothyridina sublamellosa X X Composita lewisensis X X Composita subquadrata X X Diaphragmus elegans X X Articulata Dielasma sp. X Eumetria costata X Brachiopoda Eumetria verneuliana X X Orthotetes kaskaskiensis X Punctospirifer sp. X Streptorhynchus sp. X Lingulipora sp. X Inarticulata Oehlertella pleurites X Orbiculoidea sp. X X

Table 4.10 (Continued)

Anisotrypa sp. X Archimedes cf. meekanus X X Eridopora sp. X X Fenestella sp. X X Fenestrellina sp. X Glyptopora punctipora X Lyroporella sp. X Bryozoa Stenolaemata Polypora sp. X X Rhombopora sp. X X Septopora sp. X X Sulcoretepora sp. X Thamniscus? X Unidentifiable encrusting bryozoan X Unidentifiable edrioasteroid X Edrioasteroidea Lepidodiscus laudoni X Blastoidea Pentremites elegans X X Lepidesthes formosa X Echinoidea Archaeocidaris megastylus X X Ophiuroidea Unidentifiable encrinasterid X Acrocrinus shumardi X Agassizocrinus lobatus X X Anartiocrinus maxvillensis X X Aphelecrinus mundus X X Camptocrinus cirrifer X X Cryphiocrinus girtyi X X Cymbiocrinus grandis X X Echinodermata Cymbiocrinus tumidus X X Dasciocrinus florealis X X Eupachycrinus boydii X Crinoidea Linocrinus wachsmuthi X Pentaramicrinus gracilis X X Phacelocrinus longidactylus X Phanocrinus bimagnaramus X X Phanocrinus maniformis X X Platycrinites sp. X Pterotocrinus acutus X Talarocrinus sp. X Taxocrinus whitfieldi X X Tholocrinus spinosus X X Zeacrinites magnoliaeformis X Chondrichthyes Chordata (teeth) Agassizodus X

Table 4.10 (Continued)

Cladodus X X Ctenacanthus X Deltodopsis X Deltodus X Listracanthus? X Lophodus X Mesodmodus X Psammodus? X Polyrhizodus X Taeniodus? X Venustodus X X Chondrichthyes (dermal skin Petrodus X plates)

4.4. Census Populations by Class

Table 4.11 provides a summary of occurrences at the 213 and Valley Stone quarries based on the number of invertebrates, and vertebrates, as well as on taxon levels.

In order to group the various species taxonomically, I have grouped all the data by taxanomic classes. Twelve classes from the 213 Quarry and 14 classes from the Valley

Stone Quarry will be analyzed further via statistical analyses in Chapter 5.

Table 4.11 Quick statistics for the 213 and Valley Stone quarries by taxon levels.

213 and Valley Stone Quarry Quick Statistics 213 Valley Stone Taxon Level Quarry Quarry Number of invertebrate species 43 52 Number of vertebrate genera 2 13 Number of phyla 8 8

Table 4.11 (Continued)

Number of classes 12 14 Number of echinoderm class 3 5 Number of non-echinoderm classes 9 9 Total number of species/genera 45 65

4.4.1. Species Comparison at 213 and Valley Stone Quarries

From the 213 Quarry, top-of-slab and bottom-of-slab assemblages were designated, and both represented the same lithofacies, a fine-grained calcarenite and shale. The fauna includes 15 classes for the 213 and Valley Stone quarries. Listed below are genera/species abundances for both quarries for each representative class. In fact, the

213 Quarry site contains twelve classes whereas the Valley Stone Quarry contains fourteen classes. The Valley Stone Quarry contained sixty-five identified species, and the

213 Quarry contained forty-five species. The taxa are grouped based on their respective classes throughout the chapters. Table 4.12 shows the detailed census-population summary for the 213 Quarry. The slabs of rock were analyzed for the top and bottom fauna. The loose samples were combined with top-of-slab samples. The 213 Quarry contained 3,243 individual specimens. For comparison, Table 4.13 shows the census- population summary for the Valley Stone Quarry, where 1,894 specimens were tallied.

Table 4.14 shows the census-population summary for the Valley Stone Quarry and the four continuous lithofacies: coarse-grained calcarenite, coarse-grained, argillaceous calcarenite, fine-grained calcarenite and interbedded calcilutite, and an argillaceous calcilutite

Table 4.12 Genera/species numbers by assemblage for the 213 Quarry.

213 Quarry Census Top-of- Bottom-of- Loose Grand Population Slab Slab Samples Total

Anthozoa (1) Zaphrentoides spinulosus 5 1 1 7

Scyphozoa? (1) Sphenothallus sp. 3 4 1 8

Pelecypoda (1) Sulcatopinna missouriensis 1 0 0 1

Polychaeta? (1) Crinicaminus haneyensis 1 0 0 1

Ostracoda (3) Amphissites? 1 0 0 1 Bairdia? 1 0 0 1 Coronakirkbya? 3 0 0 3 Total Ostracoda: 5 0 0 5

Articulata (6) Anthracospirifer leidyi 181 115 20 316 Cleiothyridina sublamellosa 38 16 1 55 Composita lewisensis 9 10 0 19 Composita subquadrata 33 25 7 65 Diaphragmus elegans 16 7 0 23 Eumetria verneuliana 3 0 0 3 Total Articulata: 280 173 28 481

Inarticulata (2) Oehlertella pleurites 2 0 0 2 Orbiculoidea sp. 6 2 0 8 Total Inarticulata: 8 2 0 10

Stenolaemata (9) Archimedes cf. meekanus 1415 213 110 1738 Eridopora sp. 11 5 0 16 Fenestella sp. 10 1 1 12 Glyptopora punctipora 105 24 6 135 Lyroporella sp. 0 0 1 1 Polypora sp. 82 14 3 99

Table 4.12 (Continued)

Rhombopora sp. 241 171 20 432 Septopora sp. 3 5 0 8 Sulcoretepora sp. 3 1 0 4 Total Stenolaemata: 1870 434 141 2,445

Blastoidea (1) Pentremites elegans 17 12 12 41

Echinoidea (2) Lepidesthes formosa 1 0 0 1 Archaeocidaris megastylus 0 3 0 3 (isolated spines only) Total Echinoidea: 1 3 0 4

Crinoidea (16) Acrocrinus shumardi 1 0 0 1 Agassizocrinus lobatus (juvenile) 0 0 1 1 Anartiocrinus maxvillensis 1 0 0 1 Aphelecrinus mundus 3 0 2 5 Camptocrinus cirrifer 10 1 1 12 Cryphiocrinus girtyi 0 0 2 2 Cymbiocrinus grandis 50 1 4 55 Cymbiocrinus tumidus 0 0 11 11 Dasciocrinus florealis 23 5 7 35 Pentaramicrinus gracilis 19 3 6 28 Phanocrinus bimagnaramus 20 2 7 29 Phanocrinus maniformis 3 0 1 4 Pterotocrinus acutus 9 6 1 16 Taxocrinus whitfieldi 22 2 7 31 Tholocrinus spinosus 4 0 0 4 Zeacrinites magnoliaeformis 0 0 3 3 Total Crinoidea: 165 20 53 238

Chondrichthyes (2) Cladodus 0 0 1 1 Venustodus 0 0 1 1 Total Chondrichthyes: 0 0 2 2

213 Quarry Grand Total: 3,243

Table 4.13 Genera/species numbers by assemblage for the Valley Stone Quarry.

Valley Stone Quarry Census Slab Loose Grand Population Total Samples Total

Demospongea (1) Belemnospongia fascicularis 90 143 233

Anthozoa (1) Unidentifiable tabulate coral? 1 0 1

Scyphozoa? (2) Sphenothallus sp. 26 2 28 Paraconularia missouriensis 3 5 8 Total Scyphozoa (?): 29 7 36

Pelecypoda (1) Sulcatopinna missouriensis 1 3 4

Polychaeta? (1) Crinicaminus haneyensis 2 0 2

Articulata (11) Anthracospirifer leidyi 206 66 272 Cleiothyridina sublamellosa 5 3 8 Composita lewisensis 8 0 8 Composita subquadrata 335 75 410 Diaphragmus elegans 3 23 26 Dielasma sp. 0 1 1 Eumetria costata 2 2 4 Eumetria verneuliana 5 0 5 Orthotetes kaskaskiensis 5 2 7 Punctospirifer sp. 0 7 7 Streptorhynchus sp. 1 0 1 Total Articulata: 570 179 749

Inarticulata (2) Lingulipora sp. 0 2 2

Table 4.13 (Continued)

Orbiculoidea sp. 6 0 6 Total Inarticulata: 6 2 8

Stenolaemata (10) Anisotrypa sp. 3 0 3 Archimedes cf. meekanus 14 3 17 Eridopora sp. 1 0 1 Fenestella sp. 4 0 4 Fenestrellina sp. 1 0 1 Polypora sp. 21 3 24 Rhombopora sp. 41 4 45 Septopora sp. 1 0 1 Thamniscus? 38 2 40 Unidentifiable encrusting bryozoan 4 2 6 Total Stenolaemata: 128 14 142

Edrioasteroidea (2) Unidentifiable edrioasteroid 5 1 6 Lepidodiscus laudoni 0 1 1 Total Edrioasteroidea: 5 2 7

Blastoidea (1) Pentremites elegans 60 29 89

Echinoidea (1) Archaeocidaris megastylus 33 21 54

Ophiuroidea (1) Unidentifiable encrinasterid 24 10 34

Crinoidea (18) Agassizocrinus lobatus 16 14 30 Anartiocrinus maxvillensis 16 15 31 Aphelecrinus mundus 15 10 25 Camptocrinus cirrifer 2 0 2 Cryphiocrinus girtyi 0 1 1 Cymbiocrinus grandis 8 27 35 Cymbiocrinus tumidus 0 23 23 Dasciocrinus florealis 21 67 88

Table 4.13 (Continued)

Eupachycrinus boydii 0 1 1 Linocrinus wachsmuthi 27 21 48 Pentaramicrinus gracilis 1 15 16 Phanocrinus bimagnaramus 79 62 141 Phanocrinus maniformis 1 0 1 Phacelocrinus longidactylus 1 0 1 Platycrinites sp. 0 1 1 Talarocrinus sp. 3 3 6 Taxocrinus whitfieldi 1 10 11 Tholocrinus spinosus 1 0 1 Total Crinoidea: 192 270 462

Chondrichthyes (13) Agassizodus 0 1 1 Cladodus 0 3 3 Ctenacanthus 0 1 1 Deltodopsis 0 1 1 Deltodus 0 1 1 Listracanthus? 0 1 1 Lophodus 0 1 1 Mesodmodus 0 1 1 Petrodus 0 59 59 Psammodus? 0 1 1 Polyrhizodus 0 1 1 Taeniodus? 0 1 1 Venustodus 0 1 1 Total Chondrichthyes: 0 73 73

Valley Stone Quarry Grand Total: 1,894

Table 4.14 Genera/species numbers by assemblage for the Valley Stone Quarry by lithofacies.

Fine-grained Coarse- Coarse- calcarenites Valley Stone Quarry Census grained, Argillaceous Grand grained and Population argillaceous calcilutites Total calcarenite interbedded calcarenites calcilutites Demospongea (1) Belemnospongia fascicularis 134 16 8 75 233 91

Table 4.14 (Continued)

Anthozoa (1) Unidentifiable tabulate coral? 1 0 0 0 1

Scyphozoa (?) (2) Sphenothallus sp. 4 13 9 2 28 Paraconularia missouriensis 3 0 0 5 8 Total Scyphozoa (?): 7 13 9 7 36

Pelecypoda (1) Sulcatopinna missouriensis 1 0 0 3 4

Polychaeta (?) (1) Crinicaminus haneyensis 2 0 0 0 2 . Articulata (11) Anthracospirifer leidyi 166 26 38 42 272 Cleiothyridina sublamellosa 0 2 3 3 8 Composita lewisensis 5 1 0 2 8 Composita subquadrata 63 215 25 107 410 Diaphragmus elegans 1 3 1 21 26 Dielasma sp. 0 0 0 1 1 Eumetria costata 2 0 0 2 4 Eumetria verneuliana 2 1 0 2 5 Orthotetes kaskaskiensis 2 0 0 5 7 Punctospirifer sp. 0 2 0 5 7 Streptorhynchus sp. 1 0 0 0 1 Total Articulata: 242 250 67 190 749

Inarticulata (2) Lingulipora sp. 2 0 0 0 2 Orbiculoidea sp. 1 2 3 0 6 Total Inarticulata: 3 2 3 0 8

Stenolaemata (10) Anisotrypa sp. 2 0 0 1 3 Archimedes cf. meekanus 11 6 0 0 17 Eridopora sp. 0 0 0 1 1 Fenestella sp. 0 0 1 3 4 Fenestrellina sp. 0 0 1 0 1 Polypora sp. 2 4 5 13 24 Rhombopora sp. 13 4 0 28 45 Septopora sp. 0 0 0 1 1 Thamniscus? 16 7 14 3 40 Unidentifiable encrusting bryozoan 5 1 0 0 6 Total Stenolaemata: 49 22 21 50 142

Edrioasteroidea (2) Unidentifiable edrioasteroid 6 0 0 0 6 Lepidodiscus laudoni 0 0 0 1 1 Total Edrioasteroidea: 6 0 0 1 7

Table 4.14 (Continued)

Blastoidea (1) Pentremities elegans 29 8 2 50 89

Echinoidea (1) Archaeocidaris megastylus 1 0 14 39 54

Ophiuroidea (1) Unidentifiable encrinasterid brittle star 21 2 9 2 34

Crinoidea (18) Agassizocrinus lobatus 3 5 3 19 30 Anartiocrinus maxvillensis 21 0 3 7 31 Aphelecrinus mundus 14 7 0 4 25 Camptocrinus cirrifer 0 1 1 0 2 Cryphiocrinus girtyi 0 0 0 1 1 Cymbiocrinus grandis 15 7 0 13 35 Cymbiocrinus tumidus 8 3 0 12 23 Dasciocrinus florealis 46 8 12 22 88 Eupachycrinus boydii 0 0 0 1 1 Linocrinus wachsmuthi 36 4 5 3 48 Pentaramicrinus gracilis 2 0 0 14 16 Phanocrinus bimagnaramus 68 24 26 23 141 Phanocrinus maniformis 1 0 0 0 1 Phacelocrinus longidactylus 0 1 0 0 1 Platycrinites sp. 0 0 0 1 1 Talarocrinus sp. 4 0 2 0 6 Taxocrinus whitfieldi 8 0 0 3 11 Tholocrinus spinosus 1 0 0 0 1 Total Crinoidea: 227 60 52 123 462

Chondrichthyes (13) Agassizodus 0 1 0 0 1 Cladodus 0 1 0 2 3 Ctenacanthus 0 0 0 1 1 Deltodopsis 0 0 0 1 1 Deltodus 0 0 0 1 1 Listracanthus? 0 0 0 1 1 Lophodus 0 0 0 1 1 Mesodmodus 0 0 0 1 1 Petrodus 0 0 0 59 59 Psammodus? 0 0 0 1 1 Polyrhizodus 0 0 0 1 1 Taeniodus? 0 0 0 1 1 Venustodus 0 1 0 0 1 Total Chondrichthyes: 0 3 0 70 73

Valley Stone Quarry Total: 1,894

CHAPTER 5: RELATIVE, ORDINAL AND STATISTICAL MEASURES OF

ABUNDANCE

5.1. Introduction

This chapter deals with the comparison of different suites of fossil organisms at the two different localities, the Valley Stone Quarry and the 213 Quarry. Using the definitions of Johnson (1964) and Patzkowsky and Holland (2012), a community is a set of species that coexist at a particular location and time. Hence, finding a way to compare and evaluate the nature of the communities at the two localities becomes very important.

One of the most common methods for comparing communities and lithofacies is by examining relative, ordinal and statistical measures of abundances using the metrics of richness, diversity, evenness, relative frequency, and community density. In these analyses, the term “landscapes” as used by Patzkowsky and Holland (2012), refers to an entire continuum of communities or lithofacies at a locality. The numbers of individuals used to develop these metrics are noted, tallied and listed in Chapter 4.

5.2. Relative Abundance

One of the most important methods for comparing organisms in a community are measures of relative abundance. Relative abundance was measured to compare and contrast the similarities and differences among organism communities and between the lithofacies in which they occur in the two study areas. The stratigraphic and environmental implications of a natural assemblage of species should be a more telling

comparator than any single species. For example, a high recurrence of species associations would appear to be of considerable significance (Johnson, 1964).

In order to make comparisons, the abundance of organisms is listed by class and species in the next 50 figures and accompanying tables for the 213 and Valley Stone quarries. The classes are discussed below based on their phylogenetic position. The graphs in the following figures are color-coded based on the quarry locality and what is being compared. Figures that compare organism abundances at the 213 and Valley Stone quarries are shown in dark gray; figures that compare the classes across the landscape in the Valley Stone Quarry are orange; and figures that compare the classes across the landscape in the 213 Quarry blue. The vertical scale on most of the following charts is set at 250 or 450 individuals, whereas the number of stenolaemate bryozoan individuals may be set as high as 2,100.

5.2.1. Abundance of Demospongea in the 213 and Valley Stone quarries

The first of the described classes is the Class Demospongea in the Phylum

Porifera. The only demosponge species, Belemnospongia fascicularis, was found at the

Valley Stone Quarry and is absent at the 213 Quarry. In order to make bar-graph comparisons (Figure 5.1), a “0” was used as a placeholder for those organisms that are absent at a given locality. Even though Belemnospongia fascicularis was not found at the

213 Quarry, it is shown as a “0” on the graph in Figure 5.1. In comparing the occurrence of the Class Demospongea at the Valley Stone Quarry across the four recognized lithofacies (Figure 5.2), most of the demosponge individuals occur in the coarse-grained calcarenite lithofacies.

213 & VALLEY STONE QUARRIES: CLASS DEMOSPONGEA

450 400 350 300 233 250 200 213 150 Valley Stone 100 50 0 0 Belemnospongia fascicularis

NUMBER OF INDIVIDUALS SPECIES

Figure 5.1. Number of individuals per species from the Class Demospongea in the 213 and Valley Stone quarries. The only sponge, Belemnospongia fascicularis, was not present at the 213 Quarry.

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS DEMOSPONGEA

250 134 200 75 150 100 16 8 50 0 Demospongea

CLASS NUMBER OF INDIVIDUALS OF NUMBER Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.2. The number of individuals from the Class Demospongea in each of the four lithofacies at the Valley Stone Quarry.

5.2.2. Abundance of Anthozoa in the 213 and Valley Stone quarries

Species of the Class Anthozoa in the Phylum Cnidaria are relatively rare at both quarries. One rugosan species, Zaphrentoides spinulosus, was found at the 213 Quarry, but none were present at Valley Stone Quarry (Figure 5.3). However, a poorly preserved, unidentifiable tabulate coral did occur at Valley Stone Quarry, but was absent at the 213

Quarry. In comparing the Class Anthozoa for the 213 top-of-slab and bottom-of-slab communities, Zaphrentoides spinulosus was more abundant on the top-of-slab community (Figure 5.4). In comparing the Class Anthozoa at the Valley Stone Quarry across the four recognized lithofacies (Figure 5.5), only a single poorly preserved tabulate coral was observed in the coarse-grained calcarenite lithofacies.

213 & VALLEY STONE QUARRIES: CLASS ANTHOZOA

450 400 350 300 250 200 213 150 Valley Stone 100 50 0 1 7 0 NUMBER OF INDIVIDUALS OF NUMBER 0 Unidentifiable tabulate Zaphrentoides spinulosus coral? SPECIES

Figure 5.3. Number of individuals per species from the Class Anthozoa in the 213 and Valley Stone quarries.

213 QUARRY: TOP-OF-SLAB VS. BOTTOM-OF- SLAB COMMUNITIES: CLASS ANTHOZOA

2100 1800 1500 1200 900 600 300 6 1 0 Anthozoa

CLASS NUMBER OF INDIVIDUALS OF NUMBER

Top of Slab Bottom of Slab

Figure 5.4. The number of individuals in the Class Anthozoa in the 213 Quarry, top-of- slab vs. bottom-of-slab communities.

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS ANTHOZOA

250 200 150 100 1 0 0 0 50 0 Anthozoa

NUMBER OF INDIVIDUALS OF NUMBER CLASS

Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.5. The number of individuals from the Class Anthozoa in each of the four lithofacies at the Valley Stone Quarry.

5.2.3. Abundance of Scyphozoa (?) in the 213 and Valley Stone quarries

Species of the Class Scyphozoa (?) in the Phylum Cnidaria are relatively uncommon at both localities. Sphenothallus sp., a benthic form, was found at both quarry localities. In addition, Paraconularia missouriensis, likely planktic form as adults, was found at the Valley Stone Quarry, but was not present at the 213 Quarry (Figure 5.6). In comparing the Class Scyphozoa (?) for the 213 Quarry top-of-slab and the bottom-of-slab communities, individuals of the Scyphozoan genus Sphenothallus sp. were equally abundant in each community (Figure 5.7). In comparing the Class Scyphozoa (?) at the

Valley Stone Quarry across the four recognized lithofacies (Figure 5.8), the greatest number of Scyphozoan individuals, mostly Sphenothallus sp., were found in the coarse- grained, argillaceous calcarenite lithofacies.

213 & VALLEY STONE QUARRIES: CLASS SCYPHOZOA (?)

450

400

350

300

250

200 213 150 Valley Stone 100 28 50 8 8

NUMBER OF INDIVIDUALS OF NUMBER 0 0 Spenothallus sp. Paraconularia missouriensis SPECIES

Figure 5.6. Number of individuals per species from the Class Scyphozoa (?) in the 213 and Valley Stone quarries.

213 QUARRY: TOP-OF-SLAB VS. BOTTOM-OF- SLAB COMMUNITIES: CLASS SCYPHOZOA (?)

2100 1800 1500 1200 900 600 300 4 4 0

NUMBER OF INDIVIDUALS OF NUMBER Scyphozoa? CLASS

Top of Slab Bottom of Slab

Figure 5.7. The number of individuals in the Class Scyphozoa (?) in the 213 Quarry, top- of-slab vs. bottom-of-slab communities.

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS SCYPHOZOA (?)

250 200 150 100 7 13 9 7 50 0

NUMBER OF INDIVIDUALS OF NUMBER Scyphozoa? CLASS

Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.8. The number of individuals from the Class Scyphozoa (?) in each of the four lithofacies at the Valley Stone Quarry.

100

5.2.4. Abundance of Pelecypoda in the 213 and Valley Stone quarries

The Class Pelecypoda was the only molluscan class found at either quarry. One species, Sulcatopinna missouriensis, a large semi-infaunal species, was found at both localities (Figure 5.9), and it was very sporadic at both quarries. In comparing the Class

Pelecypoda for the 213 Quarry top-of-slab and the bottom-of-slab communities, the species Sulcatopinna missouriensis was found only in the top-of-slab community (Figure

5.10). In comparing the Class Pelecypoda at the Valley Stone across the four recognized lithofacies (Figure 5.11), Sulcatopinna missouriensis occurred only in the coarse-grained calcarenite and argillaceous calcilutite facies.

213 & VALLEY STONE QUARRIES: CLASS PELECYPODA

450 400 350 300 250 200 213 150 Valley Stone 100 50 1 4 NUMBER OF INDIVIDUALS OF NUMBER 0 Sulcatopinna missouriensis SPECIES

Figure 5.9. Number of individuals per species from the Class Pelecypoda in the 213 and Valley Stone quarries.

101

213 QUARRY: TOP-OF-SLAB VS. BOTTOM-OF- SLAB COMMUNITIES: CLASS PELECYPODA

2100 1800 1500 1200 900 600 300 1 0 0

Pelecypoda NUMBER OF INDIVIDUALS OF NUMBER CLASS

Top of Slab Bottom of Slab

Figure 5.10. The number of individuals in the Class Pelecypoda in the 213 Quarry, top- of-slab vs. bottom-of-slab communities.

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS PELECYPODA

250 200 150 100 1 0 0 3 50 0 Pelecypoda

CLASS NUMBER OF INDIVIDUALS OF NUMBER

Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.11. The number of individuals from the Class Pelecypoda in each of the four lithofacies at the Valley Stone Quarry.

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5.2.5. Abundance of Polychaeta (?) in the 213 and Valley Stone quarries

Probable individuals in the Class Polychaeta (?) from the Phylum Annelida were represented by the domichnia trace fossil, Crinicaminus haneyensis, at both the 213 and

Valley Stone quarries (Figure 5.12). In comparing the Class Polychaeta (?) for the 213

Quarry top-of-slab and the bottom-of-slab communities, a single specimen of

Crinicaminus haneyensis was found only in the top-of-slab community (Figure 5.13). In comparing the Class Polychaeta (?) at the Valley Stone Quarry across the four recognized lithofacies (Figure 5.14), two individuals of Crinicaminus haneyensis were observed only in the coarse-grained, calcarenite lithofacies.

213 & VALLEY STONE QUARRIES: CLASS POLYCHAETA (?)

450

400

350

300

250

200 213

150 Valley Stone

100

50 NUMBER OF INDIVIDUALS OF NUMBER 1 2 0 Crinicaminus haneyensis SPECIES

Figure 5.12. Number of individuals per species from the Class Polychaeta (?) in the 213 and Valley Stone quarries.

103

213 QUARRY: TOP-OF-SLAB VS. BOTTOM-OF- SLAB COMMUNITIES: POLYCHAETA (?)

2100 1800 1500 1200 900 600 300 1 0 0 Polychaeta?

CLASS NUMBER OF INDIVIDUALS OF NUMBER

Top of Slab Bottom of Slab

Figure 5.13. The number of individuals in the Class Polychaeta (?) in the 213 Quarry, top-of-slab vs. bottom-of-slab communities.

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS POLYCHAETA (?)

250 200 150 100 2 0 0 0 50 0 Polychaeta?

CLASS NUMBER OF INDIVIDUALS OF NUMBER Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.14. The number of individuals from the Class Polychaeta (?) in each of the four lithofacies at the Valley Stone Quarry.

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5.2.6. Abundance of Ostracoda in the 213 and Valley Stone quarries

Three genera from the arthropod Class Ostracoda were noted at the 213 Quarry

(Figure 5.15). Because of their poor preservation, identification was only possible to the genus level. No ostracods were found in the rocks at Valley Stone Quarry, but ostracods at the 213 Quarry were noted only in the top-of-the-slab community (Figure 5.16).

213 & VALLEY STONE QUARRIES: CLASS OSTRACODA

450

400

350

300

250

200 213

150 Valley Stone

100

50 NUMBER OF INDIVIDUALS OF NUMBER 1 0 1 0 3 0 0 Amphissites? Bairdia? Coronakirkbya? SPECIES

Figure 5.15. Number of individuals per species from the Class Ostracoda in the 213 and Valley Stone quarries.

105

213 QUARRY: TOP-OF-SLAB VS. BOTTOM-OF- SLAB COMMUNITIES: CLASS OSTRACODA

2100 1800 1500 1200 900 600 300 5 0 0 Ostracoda

CLASS NUMBER OF INDIVIDUALS OF NUMBER

Top of Slab Bottom of Slab

Figure 5.16. The number of individuals in the Class Ostracoda in the 213 Quarry, top-of- slab vs. bottom-of-slab communities.

5.2.7. Abundance of Articulata in the 213 and Valley Stone quarries

Brachiopods in the Class Articulata are well-represented at both localities. For the

213 Quarry, six species are present, and 11 species were observed at the Valley Stone

Quarry (Figure 5.17). Overall, for the 213 Quarry, Anthracospirifer leidyi was the most abundant species followed by Composita subquadrata and Cleiothyridina sublamellosa

(Figure 5.17). The least abundant species at the 213 Quarry is Eumetria verneuliana. In contrast, for the Valley Stone Quarry the most abundant species was Composita subquadrata, followed by Anthracospirifer leidyi and Diaphragmus elegans. In comparing the Class Articulata for the 213 Quarry top-of-slab and the bottom-of-slab communities, the Articulata were almost twice as abundant in the top-of-slab community as in the bottom-of-slab community (Figure 5.18). In comparing articulate brachiopod species at the Valley Stone Quarry across the four recognized lithofacies (Figure 5.19),

106

brachiopod individuals were observed across all four lithofacies, but the coarse-grained calcarenite and the coarse-grained, argillaceous calcarenite lithofacies contained the most brachiopods (Figure 5.19).

213 & VALLEY STONE QUARRIES: CLASS ARTICULATA

450 410 400 350 316 300 272 250 200 150 100 55 65 23 50 8 198 26 01 04 35 07 07 01 213

0 Valley Stone NUMBER OF INDIVIDUALS OF NUMBER

SPECIES

Figure 5.17. Number of individuals per species from the Class Articulata in the 213 and Valley Stone quarries.

107

213 QUARRY: TOP-OF-SLAB VS. BOTTOM-OF- SLAB COMMUNITIES: CLASS ARTICULATA

2100 1800 1500 1200 900 600 308 173 300 0 Articulata

NUMBER OF INDIVIDUALS OF NUMBER CLASS

Top of Slab Bottom of Slab

Figure 5.18. The number of individuals in the Class Articulata in the 213 Quarry, top-of- slab vs. bottom-of-slab communities.

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS ARTICULATA 242 250 190 250 200 150 67 100 50 0

Articulata NUMBER OF INDIVIDUALS OF NUMBER CLASS

Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.19. The number of individuals from the Class Articulata in each of the four lithofacies at the Valley Stone Quarry.

108

5.2.8. Abundance of Inarticulata in the 213 and Valley Stone quarries

One unidentified species of Orbiculoidea from the brachiopod Class Inarticulata was present at both quarries (Figure 5.20). In contrast, the inarticulate genus Lingulipora sp. was found only at the Valley Stone Quarry, whereas the species Oehlertella pleurites was found only at the 213 Quarry. In general, inarticulate brachiopods are not very common in any of the quarry communities. In comparing the the numbers of inarticulate brachiopods in the 213 Quarry top-of-slab and the bottom-of-slab communities, the

Inarticulata were more abundant in the top-of-slab community (Figure 5.18). Moreover, in comparing numbers of inarticulate brachiopods at the Valley Stone Quarry across the four recognized lithofacies (Figure 5.22), individuals were observed across three of the four lithofacies.

213 & VALLEY STONE QUARRIES: CLASS INARTICULATA

450 400 350 300 250 200 213 150 100 Valley Stone 50 0 2 8 6 2 0 0 NUMBER OF INDIVIDUALS OF NUMBER Lingulipora sp. Orbiculoidea sp. Oehlertella pleurites SPECIES

Figure 5.20. Number of individuals per species from the Class Inarticulata in the 213 and Valley Stone quarries.

109

213 QUARRY: TOP-OF-SLAB VS. BOTTOM-OF- SLAB COMMUNITIES: CLASS INARTICULATA

2100 1800 1500 1200 900 600 300 8 2 0 Inarticulata CLASS

NUMBER OF INDIVIDUALS OF NUMBER Top of Slab Bottom of Slab

Figure 5.21. The number of individuals in the Class Inarticulata in the 213 Quarry, top- of-slab vs. bottom-of-slab communities.

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS INARTICULATA

250

150 3 2 3 0 50

-50 Inarticulata

CLASS NUMBER OF INDIVIDUALS OF NUMBER Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.22. The number of individuals from the Class Inarticulata in each of the four lithofacies at the Valley Stone Quarry.

110

5.2.9. Abundance of Stenolaemata in the 213 and Valley Stone quarries

The bryozoan Class Stenolaemata is well-represented at both quarry localities.

Nine bryozoan species were preserved at the 213 Quarry (Figure 5.23), and they were the most abundant faunal elements in the 213 Quarry. The most abundant species is

Archimedes cf. meekanus followed by Rhombopora sp., Glyptopora punctipora, and

Polypora sp. However, at the Valley Stone Quarry, bryozoans were not as common, and the most abundant species is Rhombopora sp. followed by Thamniscus? and Polypora sp.

Much rarer species at the 213 Quarry include Eridopora sp., Fenestella sp., Septopora sp.

Lyroporella sp., and Sulcoretepora sp. At the Valley Stone Quarry, the rarer species include Anisotrypa sp., Fenestella sp., Polypora sp., Eridopora sp., Fenestrellina sp., and

Septopora sp. In comparing the Class Stenolaemata for the 213 Quarry top-of-slab and bottom-of-slab communities, stenolaemate bryozoans were almost five times more abundant in top-of-slab community than the bottom-of-slab community (Figure 5.24). In comparing the Class Stenolaemata at the Valley Stone Quarry across the four recognized lithofacies (Figure 5.25), individuals were observed across all four lithofacies. The coarse-grained calcarenite and argillaceous calcilutite lithofacies contained the most bryozoan individuals.

111

213 & VALLEY STONE QUARRIES: CLASS STENOLAEMATA 1738 1800 1600 1400 1200 1000 800 600 432 400 135 99 200 0 3 17 16 1 12 4 0 1 0 1 0 24 45 8 1 4 0 0 40 0 6 213 0

Valley Stone NUMBER OF INDIVIDUALS OF NUMBER

112

SPECIES

Figure 5.23. Number of individuals per species in the Class Stenolaemata in the 213 and Valley Stone quarries.

213 QUARRY: TOP-OF-SLAB VS. BOTTOM- OF-SLAB COMMUNITIES: CLASS STENOLAEMATA

2011 2100 1800 1500 1200 900 434 600 300 0 Stenolaemata

NUMBER OF INDIVIDUALS OF NUMBER CLASS

Top of Slab Bottom of Slab

Figure 5.24. The number of individuals in the Class Stenolaemata in the 213 Quarry, top- of-slab vs. bottom-of-slab communities.

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS STENOLAEMATA

250 200 150 50 49 22 100 21 50 0 Stenolaemata

CLASS NUMBER OF INDIVIDUALS OF NUMBER

Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.25. The number of individuals from the Class Stenolaemata in each of the four lithofacies at the Valley Stone Quarry.

113

5.2.10. Abundance of Edrioasteroidea in the 213 and Valley Stone quarries

Five echinoderm classes are represented in the two quarry localities, including

Edrioasteroidea. Two species of edrioasteriods, an unidentifiable edrioasteroid and

Lepidodiscus laudoni, are present at the Valley Stone Quarry (Figure 5.26). However, no edrioasteroids were found at the 213 Quarry. In comparing the Class Edrioasteroidea at the Valley Stone Quarry across the four recognized lithofacies (Figure 5.27), edrioasteroids were observed in only two of the lithofacies, the coarse-grained calcarenite and argillaceous calcilutite lithofacies.

213 & VALLEY STONE QUARRIES: CLASS EDRIOASTEROIDEA

450

400

350

300

250

200 213 150 Valley Stone 100

50 6 1 NUMBER OF INDIVIDUALS OF NUMBER 0 0 0 Unidentifiable edrioasteroid Lepidodiscus laudoni SPECIES

Figure 5.26. Number of individuals per species from the Class Edrioasteroidea in the 213 and Valley Stone quarries.

114

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS EDRIOASTEROIDEA

250 200 150 100 6 0 0 1 50 0 Edrioasteroidea

CLASS NUMBER OF INDIVIDUALS OF NUMBER

Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.27. The number of individuals in the Class Edrioasteroidea in each of the four lithofacies at the Valley Stone Quarry.

5.2.11. Abundance of Blastoidea in the 213 and Valley Stone quarries

One blastoid species, Pentremites elegans, was found at both quarry localities, but it was nearly twice as abundant at the Valley Stone Quarry (Figure 5.28). In comparing the Class Blastoidea for the 213 Quarry top-of-slab and the bottom-of-slab communities,

Pentremites elegans is more than twice as abundant in the top-of-slab community than in the bottom-of-slab community (Figure 5.29). In comparing the Class Blastoidea at the

Valley Stone Quarry across the four recognized lithofacies (Figure 5.30), the species

Pentremites elegans was observed in all four of the lithofacies, but it was most abundant in the argillaceous calcilutite lithofacies.

115

213 & VALLEY STONE QUARRIES: CLASS BLASTOIDEA

450

400

350

300

250

200 213 150 Valley Stone 89 100 41

50 NUMBER OF INDIVIDUALS OF NUMBER 0 Pentremites elegans SPECIES

Figure 5.28. Number of individuals per species from the Class Blastoidea in the 213 and Valley Stone quarries.

213 QUARRY: TOP-OF-SLAB VS. BOTTOM-OF- SLAB COMMUNITIES: CLASS BLASTOIDEA

2100 1800 1500 1200 900 600 300 29 12 0 Blastoidea

NUMBER OF INDIVIDUALS OF NUMBER CLASS

Top of Slab Bottom of Slab

Figure 5.29. The number of individuals in the Class Blastoidea in the 213 Quarry, top-of- slab vs. bottom-of-slab communities.

116

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS BLASTOIDEA

250 200 150 50 29 100 8 2 50 0 Blastoidea

NUMBER OF INDIVIDUALS OF NUMBER CLASS

Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.30. The number of individuals from the Class Blastoidea in each of the four lithofacies at the Valley Stone Quarry.

5.2.12. Abundance of Echinoidea in the 213 and Valley Stone quarries

Two echinoid species were found at the 213 Quarry locality; however, one species Archaeocidaris megastylus, was only recognized based on its isolated spines

(Figure 5.31). At the Valley Stone Quarry, Archaeocidaris megastylus is far more abundant with 54 individuals recorded. In comparing the Class Echinoidea for the 213 top-of-slab and bottom-of-slab communities, more echinoids were recognized in the bottom-of-slab community (Figure 5.32). In comparing the Class Echinoidea at the

Valley Stone Quarry across the four recognized lithofacies (Figure 5.33), complete

Archaeocidaris megastylus individuals were observed in three of the four of the lithofacies. The argillaceous-calcilutite lithofacies contained the most individuals.

117

213 & VALLEY STONE QUARRIES: CLASS ENCHINOIDEA

450 400 350 300 250 200 213 150 Valley Stone 100 54 50 3 1 0 0

Archaeocidaris megastylus Lepidesthes formosa NUMBER OF INDIVIDUALS OF NUMBER SPECIES

Figure 5.31. Number of individuals per species from the Class Echinoidea in the 213 and Valley Stone quarries.

213 QUARRY: TOP-OF-SLAB VS. BOTTOM-OF- SLAB COMMUNITIES: CLASS ECHINOIDEA

2100 1800 1500 1200 900 600 300 1 3 0

Echinoidea NUMBER OF INDIVIDUALS OF NUMBER CLASS

Top of Slab Bottom of Slab

Figure 5.32. The number of individuals in the Class Echinoidea in the 213 Quarry, top- of-slab vs. bottom-of-slab communities.

118

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS ECHINOIDEA

250 200 150 39 100 1 0 14 50 0 Echinoidea

CLASS NUMBER OF INDIVIDUALS OF NUMBER

Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.33. The number of individuals from the Class Echinoidea in each of the four lithofacies at the Valley Stone Quarry.

5.2.13. Abundance of Ophiuroidea in the 213 and Valley Stone quarries

Ophiuroids, or brittle stars, are only present at the Valley Stone Quarry locality and are represented by a new species of encrinasterid brittle star (Figure 5.34). In comparing the Class Ophiuroidea at the Valley Stone Quarry across the four recognized lithofacies (Figure 5.35), individuals were observed in all four of the lithofacies, but they are most abundant in the coarse-grained calcarenite lithofacies.

119

213 & VALLEY STONE QUARRIES: CLASS OPHIUROIDEA

450 400 350 300 250 200 213 150 Valley Stone 100 34 50 0

NUMBER OF INDIVIDUALS OF NUMBER 0 Unidentifiable encrinasterid brittle star SPECIES

Figure 5.34. The number of individuals in the Class Ophiuroidea in the 213 and Valley Stone quarries.

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS OPHIUROIDEA

250 200 150 21 100 2 9 2 50 0 Ophiuroidea

CLASS NUMBER OF INDIVIDUALS OF NUMBER

Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.35. The number of individuals from the Class Ophiuroidea in each of the four lithofacies at the Valley Stone Quarry. 120

5.2.14. Abundance of Crinoidea in the 213 and Valley Stone quarries

Of all the echinoderm classes, the Crinoidea are the most abundant in terms of the number of species represented at each locality with 21 species recorded total (Figure

5.36). For the 213 Quarry, sixteen crinoid species are present, with the most abundant being Cymbiocrinus grandis, Dasciocrinus florealis, and Taxocrinus whitfieldi. For the

Valley Stone Quarry, eighteen species are present; the most abundant include

Phanocrinus bimagnaramus, Dasciocrinus florealis, and Linocrinus wachsmuthi. The least abundant at the 213 Quarry are the Acrocrinus shumardi, Agassizocrinus lobatus, and Anartiocrinus maxvillensis, each with one individual. For the Valley Stone Quarry, the least abundant are the Cryphiocrinus girtyi, Eupachycrinus boydii, Phanocrinus maniformis, Phacelocrinus longidactylus, Platycrinites sp., Tholocrinus spinosus, each also with one individual. In comparing the Class Crinoidea for the 213 Quarry top-of-slab and the bottom-of-slab communities, crinoids are nearly 11 times more abundant for the top-of-slab community than in the bottom-of-slab community (Figure 5.37). In comparing the Class Crinoidea at the Valley Stone Quarry across the four recognized lithofacies (Figure 5.38), crinoid individuals were observed in all four of the lithofacies, but they are most abundant in the coarse-grained-calcarenite lithofacies.

The crinoids can be broken down further into their respective subclasses. The crinoid Subclass Cladida predominates across all lithofacies and communities at the two quarry localities (Figure 5.39). In the Valley Stone Quarry, cladids predominate in all lithofacies, but they are most abundant in the coarse-grained calcarenite lithofacies

(Figure 5.40).

121

213 & VALLEY STONE QUARRIES: CLASS CRINOIDEA

450

400

350

300

250

200

122 141 150 213

100 88 Valley Stone NUMBER OF INDIVIDUALS OF NUMBER

55 48 50 31 35 35 31 30 25 23 28 29 12 11 16 16 11 1 0 1 1 5 2 2 1 0 1 0 4 1 0 1 0 1 0 0 6 4 1 3 0 0

SPECIES

Figure 5.36. Number of individuals per species from the Class Crinoidea in the 213 and Valley Stone quarries.

213 QUARRY: TOP-OF-SLAB VS. BOTTOM- OF-SLAB COMMUNITIES: CLASS CRINOIDEA

2100 1800 1500 1200 900 600 218 300 20 0 Crinoidea

CLASS NUMBR NUMBR INDIVIDUALS OF

Top of Slab Bottom of Slab

Figure 5.37. The number of individuals in the Class Crinoidea in the 213 Quarry, top-of- slab vs. bottom-of-slab communities.

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS CRINOIDEA

227 250 200 123 150 60 52 100 50 0 Crinoidea

NUMBER OF INDIVIDUALS OF NUMBER CLASS

Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.38. The number of individuals from the Class Crinoidea in each of the four lithofacies at the Valley Stone Quarry.

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213 Quarry - Crinoidea by Subclass and Lithofacies 200 178 167 150

100

50 22 29 29 31 7 11 2 0

NUMBER OF INDIVIDUALS OF NUMBER CAMERATA CLADIDA FLEXIBILIA SUBCLASS

Top of Slab Bottom of Slab 213 Total

Figure 5.39. The Crinoids by subclass for the 213 Quarry locality, top-of-slab and bottom-of-slab communities.

Valley Stone Quarry - Crinoidea by Subclass and Lithofacies 442 450 400 350 300 250 215 200 150 119 100 59 49

50 4 1 3 1 9 8 0 0 3 11 NUMBER OF INDIVIDUALS OF NUMBER 0 CAMERATA CLADIDA FLEXIBILIA SUBCLASS

Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenites Fine-grained calcarenites and interbedded calcilutites Argillaceous calcilutites Valley Stone Total

Figure 5.40. Crinoids by subclass across the entire Valley Stone Quarry “landscape”: Coarse-grained calcarenite, coarse-grained argillaceous calcarenite, fine-grained calcarenite and interbedded calcilutite, and argillaceous calcilutite lithofacies.

124

5.2.15. Abundance of Chondrichthyes in the 213 and Valley Stone quarries

The Class Chondrichthyes is a vertebrate, cartilaginous class of fish that includes the sharks and rays. Typically, their only hard parts include dermal skin plates and teeth, which occur locally at both quarry localities. As these teeth and plates probably had several forms in any one individual, the genus names used are merely “form genera.”

Their presence in the lithofacies and communities indicates that these fish were probably swimming above and within the communities, possibly preying on some of the invertebrate species.

The Class Chondrichthyes is especially well-represented at the Valley Stone

Quarry, with thirteen form genera (Figure 5.41), whereas, only two genera are present in the 213 Quarry communities. Of all of the form genera, the most abundant was the

Petrodus, representing several lenses of dermal skin plates from the Valley Stone Quarry.

At the 213 Quarry locality, chondrichthyan fossils only occur in the top-of-slab community (Figure 5.42). In comparing the Class Chondrichthyes at the Valley Stone

Quarry across the four recognized lithofacies (Figure 5.43), chondrichthyan fossils are only present in two of the four of the lithofacies, and most of them occur in the argillaceous-calcilutite lithofacies.

125

213 & VALLEY STONE QUARRIES: CLASS CHONDRICHTHYES

450 400 350 300 250 200 150 213 100 59 Valley Stone 50 01 13 01 01 01 01 01 01 0 01 01 01 11

0 NUMBER OF INDIVIDUALS OF NUMBER

SPECIES

Figure 5.41. Number of form genera from the Class Chondrichthyes in the 213 and

Valley Stone quarries.

213 QUARRY: TOP-OF-SLAB VS. BOTTOM- OF-SLAB COMMUNITIES: CLASS CHONDRICHTHYES

2100 1800 1500 1200 900 600 300 2 0 0 Chondrichthyes

CLASS NUMBER OF INDIVIDUALS OF NUMBER

Top of Slab Bottom of Slab

Figure 5.42. The number of individuals in the Class Chondrichthyes in the 213 Quarry, top-of-slab vs. bottom-of-slab communities.

126

VALLEY STONE, ENTIRE "LANDSCAPE": CLASS CONDRICHTHYES

250 200 150 70 100 0 3 0 50 0 Chondrichthyes

CLASS NUMBER OF INDIVIDUALS OF NUMBER

Coarse-grained, calcarenite Coarse-grained, argillaceous calcarenite Fine-grained, calcarenite and interbedded calcilutite Argillaceous calcilutite

Figure 5.43. The number of specimens in the Class Chondrichthyes in each of the four lithofacies at the Valley Stone Quarry.

5.3. 213 and Valley Stone quarries: Comparison of numerical abundances by class

The two studied localities can be broken down further by community and lithofacies, which probably represent distinct depositional environments. Each environment will be analyzed based on the abundance of individuals in each class.

Overall, the 213 and Valley Stone quarries are compared by class in Figure 5.44. The

Class Stenolaemata (Bryozoa) is overall most abundant in the 213 Quarry communities, followed by the Class Articulata (Brachiopoda) and the Class Crinoidea (Echinodermata).

For the Valley Stone Quarry, the Class Articulata (Brachiopoda) is the most abundant class, followed by the classes Crinoidea and Demospongea (Figure 5.44).

127

213 & Valley Stone Quarries Abundance by Class 2445 2500

2000

1500

1000 749 481 462 500 233 238 142 89

0 7 1 8 36 1 4 1 2 5 0 10 8 0 7 41 4 54 0 41 273 NUMBER OF INDIVIDUALS OF NUMBER 0

CLASS

213 Valley Stone

Figure 5.44. Comparison of numerical abundances by class for the 213 and Valley Stone quarries.

5.4. 213 Quarry Community Abundance by Class

Both communities together at the 213 Quarry contain slightly more than 3,200 individuals of all species. However, at almost 2,600 individuals (Figure 5.45), the top-of- slab community exhibits almost four times the number of individuals as does the bottom- of-slab community (649; Figure 5.46). More detailed comparisons are provided in the next section.

5.4.1. The 213 Quarry: Top-of-slab vs. bottom-of-slab abundances by Class

Both the top-of-slab and bottom-of-slab communities at the 213 Quarry are dominated by bryozoans (Class Stenolaemata) particularly A. cf. meekanus, followed in

128

abundance at both communities by articulate brachiopods, especially A. leidyi and C. subquadrata (Figures 5.45 and 5.46). Crinoids come in a distant third in both communities but are relatively uncommon in the bottom-of-slab community. Bryozoans are nearly five times more abundant; brachiopods almost twice as abundant; and crinoids nearly 11 times more abundant in the top-of-slab community than the same classes are in the bottom-of-slab community.

213 Quarry: Top-of-slab Community, Abundance by Class

2500 2011 Anthozoa 2000 Scyphozoa? 1500 Pelecypoda 1000 Polychaeta? 308 500 218 Ostracoda 6 4 1 1 5 8 29 1 2 Articulata

0 Inarticulata

Stenolaemata NUMBER OF INDIVIDUALS OF NUMBER Blastoidea CLASS Echinoidea

Figure 5.45. Number of individuals by class in the top-of-slab community at the 213 Quarry locality. 2,594 individuals are present.

129

213 Quarry: Bottom-of-slab Community, Abundance by Class

2500 Anthozoa

2000 Scyphozoa? Pelecypoda 1500 Polychaeta? 1000 Ostracoda 434 500 173 Articulata 1 4 0 0 0 2 12 3 20 0 0 Inarticulata

NUMBER OF INDIVIDUALS OF NUMBER Stenolaemata Blastoidea Echinoidea CLASS Crinoidea

Figure 5.46. Number of individuals by class for the bottom-of-slab community at the 213 Quarry locality. 649 individuals are present.

5.5. Valley Stone Quarry Lithofacies Abundance by Class

In the Valley Stone Quarry, faunas were recorded relative to the four contiguous lithofacies. Nearly 1,900 individuals were recorded across the four Valley Stone Quarry lithofacies, and these faunas are compared by lithofacies in the next section. As the faunas in each lithofacies represent a group of organisms at a given time and place, by the definitions of Johnson (1964) and Patzkowsky and Holland (2012), they are communities. Henceforth, I will deal with them as communities.

5.5.1. Lithofacies Communities: Abundance by Class

Both the coarse-grained calcarenite and the coarse-grained, argillaceous calcarenite communities at the Valley Stone Quarry are dominated by brachiopods (Class

Articulata) particularly C. subquadrata, followed in abundance in both communities by

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the crinoids, especially P. bimagnaramus and D. florealis. The sponges (Class

Demospongea) come in a distant third in the coarse-grained, calcarenite, community but are relatively uncommon in the adjacent coarse-grained, argillaceous calcarenite community (Figures 5.47 and 5.48).

Overall, the nearly 723 individuals in the coarse-grained, calcarenite community are nearly two times more populous than those in the coarse-grained, argillaceous calcarenite community. Crinoids are nearly four times more abundant, brachiopods equally abundant, and bryozoans nearly two times more abundant in the coarse-grained, calcarenite community than the same classes are in the coarse-grained, argillaceous calcarenite community (Figures 5.47 and 5.48).

Brachiopods (Class Articulata), particularly C. subquadrata, followed in both

communities by the crinoids, especially P. bimagnaramus, predominate in both the fine- grained calcarenite and interbedded calcilutite and the argillaceous calcilutite communities at the Valley Stone Quarry. The sponges (Class Demospongea) come in a distant third in the argillaceous calcilutite community, but are relatively uncommon in fine-grained calcarenite community (Figures 5.49 and 5.50).

Overall, the nearly 610 individuals in the argillaceous calcilutite community are nearly five times more populous than those in the fine-grained calcarenite and interbedded calcilutite community. Crinoids are nearly three times more abundant, brachiopods are three times more abundant, and bryozoans nearly two times more abundant in the argillaceous calcilutite community than the same classes are in the fine- grained calcarenite and interbedded calcilutite community (Figures 5.49 and 5.50).

131

Valley Stone Quarry: Coarse-grained, Calcarenite Community, Abundance by Class 242 227 250 Demospongea 200 Anthozoa 134 150 Scyphozoa? 100 Pelecypoda 49 50 29 21 Polychaeta? 1 7 1 2 3 6 1 0 0 Articulata Inarticulata

Stenolaemata NUMBER OF INDIVIDUALS OF NUMBER Edrioasteroidea CLASS Blastoidea

Figure 5.47. Number of individuals by class for the coarse-grained, calcarenite community at the Valley Stone Quarry locality. 723 individuals are present.

Valley Stone Quarry: Coarse-grained, Argillaceous Calcarenite Community, Abundance by Class 250 250 Demospongea 200 Anthozoa 150 Scyphozoa? 100 60 22 Pelecypoda 50 16 0 13 0 0 2 0 8 0 2 3 0 Polychaeta? Articulata Inarticulata

NUMBER OF INDIVIDUALS OF NUMBER Stenolaemata CLASS Edrioasteroidea

Figure 5.48. Number of individuals by class for the coarse-grained, argillaceous calcarenite community at the Valley Stone Quarry locality. 376 individuals are present.

132

Valley Stone Quarry: Fine-grained Calcarenite and Interbedded Calcilutite Community, Abundance by Class

250 Demospongea 200 Anthozoa 150 Scyphozoa? 100 67 52 Pelecypoda 50 21 Polychaeta? 8 0 9 0 0 3 0 2 14 9 0 0 Articulata Inarticulata

NUMBER OFINDIVIDUALS NUMBER Stenolaemata Edrioasteroidea CLASS Blastoidea

Figure 5.49. Number of individuals by class for the fine-grained calcarenite and interbedded calcilutite community at the Valley Stone Quarry locality. 185 individuals are present.

Valley Stone Quarry: Argillaceous-Calcilutite Community, Abundance by Class

250 Demospongea 190 Anthozoa 200 Scyphozoa? 150 123 Pelecypoda

100 75 70 Polychaeta? 50 50 39 Articulata 50 0 7 3 0 0 1 2 Inarticulata 0 Stenolaemata NUMBER OFINDIVIDUALS NUMBER Edrioasteroidea Blastoidea Echinoidea CLASS Ophiuroidea

Figure 5.50. Number of individuals by class for the argillaceous calcilutite community at the Valley Stone Quarry locality. 610 individuals are present.

133 5.6. Ordinal Estimates of Relative Abundance

In order to compare the organisms found at the 213 and Valley Stone quarries and the various lithofacies communities, multiple measurements of diversity were used to capture the different aspects of individual communities and the landscapes as a whole. To compare the fossil organisms and communities at each quarry, I analyzed the census- population data (Appendix A) for richness, diversity, and evenness, using several statistical methods. The data used for developing the measures of richness, diversity, and evenness were analyzed using census-population counts from the top- and bottom-of-slab communities from the 213 Quarry (Figures 5.45 and 5.46; Appendix A), as well as for collections from the four lithofacies communities at the Valley Stone Quarry (Figures

5.47 through 5.50; Appendix A). Descriptive interpretations of abundance vary from author to author, but the interpretations used in Table 5.1 were modified from the ecological work of Adler (2010) and Brett et al. (2007). Table 5.1 was modified to categorically describe each species based on its relative frequency. Seven descriptors were used to describe the relative frequencies based on percentages: absent, sporadic, rare, occasional, common, frequent, and dominant (Table 5.1).

Abundance descriptors were assigned to each species in the 213 Quarry top-of- slab, bottom-of-slab and whole quarry landscape communities (Table 5.2). Similar descriptors were assigned to each species from Valley Stone Quarry lithofacies, communities and the community landscape as a whole (Table 5.3).

134 Table 5.1 Species Abundance Descriptors and Percentages.

Species Abundance Percentage x (Relative Abundance Description Frequency as %) Absent x = 0% Sporadic 0% < x ≤ 1% Rare 1% < x ≤ 5% Occasional 5% < x ≤ 10% Common 10% < x ≤ 30% Frequent 30% < x ≤ 50% Dominant 50% < x ≤ 100%

5.6.1. Ordinal Estimates of Relative Abundance for 213 Quarry (Table 5.2)

In order to understand 213 Quarry relative-frequency data, species abundance descriptors were assigned to each species based on Table 5.1. For the top-of-slab community, the majority of species fell into the sporadic or rare category (Table 5.2). The brachiopod A. leidyi fell into the occasional category. The small ramose bryozoan,

Rhombopora sp., was listed as common, whereas the fenestrate bryozoan, A. cf. meekanus was the dominant organism in the community. No species fell into the frequent category.

For the bottom-of-slab community, the majority of species fell into the sporadic and the rare categories. Rhombopora sp. and A. leidyi were listed as common, whereas the occurrence of A. cf. meekanus was listed as frequent. No species were listed in the occasional and dominant categories.

For the 213 landscape as a whole, the majority of species (42) fell into the sporadic and rare categories. A. leidyi fell into the occasional category. Rhombopora sp. 135 was listed as common. The bryozoan, A. cf. meekanus was clearly the dominant species in the community.

Forty-five species were observed across the 213 Quarry “landscape,” but the vast majority of those species (42) were only sporadic or rare in occurrence (Table 5.2). Only three species were abundant enough to be classified in the occasional-to-dominant categories, and these include the articulate brachiopod, A. leidyi, as occasional, the stenolaemate bryozoan, Rhombopora sp., as common, and the stenolaemate bryozoan A. cf. meekanus, as dominant.

Moreover, the total abundance of these three species is nearly 77% in both top-of- slab and bottom-of-slab communities, and the rank of the species remains the same in both communities. However, the percentage of A. cf. meekanus in the top-of-slab community is almost double that of the bottom-of-slab community. In contrast, the percentage of Rhombopora sp. and A. leidyi in the bottom-of-slab community is nearly double that of those same species in the top-of-slab community.

Table 5.2 The relative frequency of species as a percentage and abundance descriptors for each community and the entire landscape at the 213 Quarry.

213 213 Top- 213 213 Entire 213 Entire 213 Top- Bottom-of- of-Slab Bottom- Landscape Landscape of-Slab Slab of-Slab Relative 213 Quarry Taxa Species Species Relative Species Abundance Relative Frequency Abundance Frequency Abundance Description Frequency Total Description Description Anthozoa Zaphrentoides spinulosus 0.23% Sporadic 0.15% Sporadic 0.22% Sporadic

Schyphozoa?

136 Table 5.2 (Continued)

Sphenothallus sp. 0.15% Sporadic 0.62% Sporadic 0.25% Sporadic

Pelecypoda Sulcatopinna missouriensis 0.04% Sporadic 0.00% Absent 0.03% Sporadic

Polychaeta? Crinicaminus haneyensis 0.04% Sporadic 0.00% Absent 0.03% Sporadic

Ostracoda Amphissites? 0.04% Sporadic 0.00% Absent 0.03% Sporadic Bairdia? 0.04% Sporadic 0.00% Absent 0.03% Sporadic Coronakirkbya? 0.12% Sporadic 0.00% Absent 0.09% Sporadic

Articulata Anthracospirifer leidyi 7.75% Occasional 17.72% Common 9.74% Occasional Cleiothyridina sublamellosa 1.50% Rare 2.47% Rare 1.70% Rare Composita lewisensis 0.35% Sporadic 1.54% Rare 0.59% Sporadic Composita subquadrata 1.54% Rare 3.85% Rare 2.00% Rare Diaphragmus elegans 0.62% Sporadic 1.08% Rare 0.71% Sporadic Eumetria verneuliana 0.12% Sporadic 0.00% Absent 0.09% Sporadic

Inarticulata Oehlertella pleurites 0.08% Sporadic 0.00% Absent 0.06% Sporadic Orbiculoidea sp. 0.23% Sporadic 0.31% Sporadic 0.25% Sporadic

Stenolaemata Archimedes cf. meekanus 58.79% Dominant 32.82% Frequent 53.59% Dominant Eridopora sp. 0.42% Sporadic 0.77% Sporadic 0.49% Sporadic Fenestella sp. 0.42% Sporadic 0.15% Sporadic 0.37% Sporadic Glyptopora punctipora 4.28% Rare 3.70% Rare 4.16% Rare Lyroporella sp. 0.04% Sporadic 0.00% Absent 0.03% Sporadic Polypora sp. 3.28% Rare 2.16% Rare 3.05% Rare Rhombopora sp. 10.06% Common 26.35% Common 13.32% Common Septopora sp. 0.12% Sporadic 0.77% Sporadic 0.25% Sporadic Sulcoretepora sp. 0.12% Sporadic 0.15% Sporadic 0.12% Sporadic

Blastoidea

137 Table 5.2 (Continued)

Pentremites elegans 1.12% Rare 1.85% Rare 1.26% Rare

Echinoidea Lepidesthes formosa 0.04% Sporadic 0.00% Absent 0.03% Sporadic Archaeocidaris megastylus 0.00% Absent 0.46% Sporadic 0.09% Sporadic (isolated spines only)

Crinoidea Acrocrinus shumardi 0.04% Sporadic 0.00% Absent 0.03% Sporadic Agassizocrinus lobatus (juvenile) 0.04% Sporadic 0.00% Absent 0.03% Sporadic Anartiocrinus maxvillensis 0.04% Sporadic 0.00% Absent 0.03% Sporadic Aphelecrinus mundus 0.19% Sporadic 0.00% Absent 0.15% Sporadic Camptocrinus cirrifer 0.42% Sporadic 0.15% Sporadic 0.37% Sporadic Cryphiocrinus girtyi 0.08% Sporadic 0.00% Absent 0.06% Sporadic Cymbiocrinus grandis 2.08% Rare 0.15% Sporadic 1.70% Rare Cymbiocrinus tumidus 0.42% Sporadic 0.00% Absent 0.34% Sporadic Dasciocrinus florealis 1.16% Rare 0.77% Sporadic 1.08% Rare Pentaramicrinus gracilis 0.96% Sporadic 0.46% Sporadic 0.86% Sporadic Phanocrinus bimagnaramus 1.04% Rare 0.31% Sporadic 0.89% Sporadic Phanocrinus maniformis 0.15% Sporadic 0.00% Absent 0.12% Sporadic Pterotocrinus acutus 0.39% Sporadic 0.92% Sporadic 0.49% Sporadic Taxocrinus whitfieldi 1.12% Rare 0.31% Sporadic 0.96% Sporadic Tholocrinus spinosus 0.15% Sporadic 0.00% Absent 0.12% Sporadic Zeacrinites magnoliaeformis 0.12% Sporadic 0.00% Absent 0.09% Sporadic

Chondrichthyes Cladodus 0.04% Sporadic 0.00% Absent 0.03% Sporadic Venustodus 0.04% Sporadic 0.00% Absent 0.03% Sporadic

Grand Total: 100.00% 100.00% 100.00%

138 5.6.2. Ordinal Estimates of Relative Abundance for the Valley Stone Quarry

In order to understand the relative species abundance at the Valley Stone Quarry, species-abundance descriptors (Table 5.1) were assigned based on the percentages in

Table 5.3. Of the 65 species collected at the Valley Stone Quarry, all but four species, including the demosponge, B. fascicularis, the brachiopods A. leidyi and C. subquadrata, and the crinoid P. bimagnaramus, were classified as “rare” or “sporadic.” Species descriptors for categories higher than rare, vary from one lithofacies to another.

Of the 41 species found in the coarse-grained calcarenite community, only five species occur in abundances great enough to be classified higher than rare. The demosponge B. fascicularis and the brachiopod A. leidyi exhibit the highest numbers of individuals, comprise about 42% of the fauna, and are classified as “common.” The brachiopod C. subquadrata and the crinoid P. bimagnaramus comprise about 24% of the fauna and are classfied as occasional. The coarse-grained agrillacaeous calcarenite community contains 28 species, and except for C. subquadrata, which is the “dominant” form, and A. leidyi and P. bimagnaramus, which are “occasional,” all other species are rare or sporadic.

In the fine-grained calcarenite and interbedded calcilutite community, which contains 21 species, the brachiopods, A. leidyi and C. subquadrata, with a total of about

35% were classified as “common.” The bryozoan Thamniscus (?), the echinoid A. megastylus, an unidentified ophiuroid, and the crinoid D. florealis were classified as occasional. All other species are rare or sporadic.

139

In contrast, the argillaceous calcilutite community contains 49 species, the greatest number of any community at the Valley Stone Quarry. Of these species, the brachiopod C. subquadrata was most abundant at 17.5%, followed by the sponge B. fascicularis at 12.3%. These were the only “common” species and together comprise about 30% of the fauna. Of the other faunal elements, the brachiopod A. leidyi, the blastoid P. elegans, the echinoid A. megastylus, and chondrichthyes plates are classified as “occasional,” and reflect about 33% of the fauna. Expect for these six species, the other 43 species in the community are classified as “rare” or “sporadic.”

Although collections from this quarry were made because of the echinoderms, the abundance of echinoderms never rises above the “occasional” level. Other classes, in this case brachiopods or demosponges, are far more abundant.

140

Table 5.3 The relative frequency as a percentage and abundance descriptors for the Valley Stone Quarry lithofacies communities and landscape.

Fine- Coarse- Fine- Corase- Coarse- grained, Coarse- grained, grained, Arg. VS Entire VS Entire grained, grained, calcarenites Arg. grained, arg. calcarenites calcilutites Landscape Landscape Valley Stone (VS) Quarry calcarenite arg. and intbd calcilutites calcarenite calcarenite and intbd Species Relative Species Taxa Species calcarenite calcilutite Relative Relative Species calcilutite Abund. Frequency Abund. Abund. Relative Species Frequency Frequency Abund. Relative Descr. Total Descr. Descr. Frequency Abund. Descr. Frequency Descr.

Demospongea Belemnospongia fascicularis 18.53% Common 4.26% Rare 4.32% Rare 12.30% Common 12.26% Common

Anthozoa

141 Unidentifiable tabulate coral? 0.14% Sporadic 0.00% Absent 0.00% Absent 0.00% Absent 0.05% Sporadic

Scyphozoa? Sphenothallus sp. 0.55% Sporadic 3.46% Rare 4.86% Rare 0.33% Sporadic 1.47% Rare Paraconularia missouriensis 0.41% Sporadic 0.00% Absent 0.00% Absent 0.82% Sporadic 0.42% Sporadic

Pelecypoda Sulcatopinna missouriensis 0.14% Sporadic 0.00% Absent 0.00% Absent 0.49% Sporadic 0.21% Sporadic

Polychaeta? Crinicaminus haneyensis 0.28% Sporadic 0.00% Absent 0.00% Absent 0.00% Absent 0.11% Sporadic

Articulata Anthracospirifer leidyi 22.96% Common 6.91% Occasional 20.54% Common 6.89% Occasional 14.31% Common Cleiothyridina sublamellosa 0.00% Absent 0.53% Sporadic 1.62% Rare 0.49% Sporadic 0.42% Sporadic Composita lewisensis 0.69% Sporadic 0.27% Sporadic 0.00% Absent 0.33% Sporadic 0.42% Sporadic

Table 5.3 (Continued)

Composita subquadrata 8.71% Occasional 57.18% Dominant 13.51% Common 17.54% Common 21.57% Common Diaphragmus elegans 0.14% Sporadic 0.80% Sporadic 0.54% Sporadic 3.44% Rare 1.37% Rare Dielasma sp. 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Eumetria costata 0.28% Sporadic 0.00% Absent 0.00% Absent 0.33% Sporadic 0.21% Sporadic Eumetria verneuliana 0.28% Sporadic 0.27% Sporadic 0.00% Absent 0.33% Sporadic 0.26% Sporadic Orthotetes kaskaskiensis 0.28% Sporadic 0.00% Absent 0.00% Absent 0.82% Sporadic 0.37% Sporadic Punctospirifer sp. 0.00% Absent 0.53% Sporadic 0.00% Absent 0.82% Sporadic 0.37% Sporadic Streptorhynchus sp. 0.14% Sporadic 0.00% Absent 0.00% Absent 0.00% Absent 0.05% Sporadic

Inarticulata Lingulipora sp. 0.28% Sporadic 0.00% Absent 0.00% Absent 0.00% Absent 0.11% Sporadic Orbiculoidea sp. 0.14% Sporadic 0.53% Sporadic 1.62% Rare 0.00% Absent 0.32% Sporadic

142 Stenolaemata

Anisotrypa sp. 0.28% Sporadic 0.00% Absent 0.00% Absent 0.16% Sporadic 0.16% Sporadic Archimedes cf. meekanus 1.52% Rare 1.60% Rare 0.00% Absent 0.00% Absent 0.89% Sporadic Eridopora sp. 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Fenestella sp. 0.00% Absent 0.00% Absent 0.54% Sporadic 0.49% Sporadic 0.21% Sporadic Fenestrellina sp. 0.00% Absent 0.00% Absent 0.54% Sporadic 0.00% Absent 0.05% Sporadic Polypora sp. 0.28% Sporadic 1.06% Rare 2.70% Rare 2.13% Rare 1.26% Rare Rhombopora sp. 1.80% Rare 1.06% Rare 0.00% Absent 4.59% Rare 2.37% Rare Septopora sp. 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Thamniscus? 2.21% Rare 1.86% Rare 7.57% Occasional 0.49% Sporadic 2.10% Rare

Unidentifiable encrusting bryozoan 0.69% Sporadic 0.27% Sporadic 0.00% Absent 0.00% Absent 0.32% Sporadic

Edrioasteroidea Unidentifiable edrioasteroid 0.83% Sporadic 0.00% Absent 0.00% Absent 0.00% Absent 0.32% Sporadic Lepidodiscus laudoni 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic

Table 5.3 (Continued)

Blastoidea Pentremites elegans 4.01% Rare 2.13% Rare 1.08% Rare 8.20% Occasional 4.68% Rare

Echinoidea Archaeocidaris megastylus 0.14% Sporadic 0.00% Absent 7.57% Occasional 6.39% Occasional 2.84% Rare

Ophiuroidea Unidentifiable encrinasterid brittle star 2.90% Rare 0.53% Sporadic 4.86% Occasional 0.33% Sporadic 2.16% Rare

Crinoidea Agassizocrinus lobatus 0.41% Sporadic 1.33% Rare 1.62% Rare 3.11% Rare 1.58% Rare Anartiocrinus maxvillensis 2.90% Rare 0.00% Absent 1.62% Rare 1.15% Rare 1.63% Rare

143 Aphelecrinus mundus 1.94% Rare 1.86% Rare 0.00% Absent 0.66% Sporadic 1.32% Rare Camptocrinus cirrifer 0.00% Absent 0.27% Sporadic 0.54% Sporadic 0.00% Absent 0.11% Sporadic

Cryphiocrinus girtyi 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Cymbiocrinus grandis 2.07% Rare 1.86% Rare 0.00% Absent 2.13% Rare 1.84% Rare Cymbiocrinus tumidus 1.11% Rare 0.80% Sporadic 0.00% Absent 1.97% Rare 1.21% Rare Dasciocrinus florealis 6.36% Occasional 2.13% Rare 6.49% Occasional 3.61% Rare 4.63% Rare Eupachycrinus boydii 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Linocrinus wachsmuthi 4.98% Rare 1.06% Rare 2.70% Rare 0.49% Sporadic 2.52% Rare Pentaramicrinus gracilis 0.28% Rare 0.00% Absent 0.00% Absent 2.30% Rare 0.84% Sporadic Phanocrinus bimagnaramus 9.41% Occasional 6.38% Occasional 14.05% Common 3.77% Rare 7.42% Occasional Phanocrinus maniformis 0.14% Sporadic 0.00% Absent 0.00% Absent 0.00% Absent 0.05% Sporadic Phacelocrinus longidactylus 0.00% Absent 0.27% Sporadic 0.00% Absent 0.00% Absent 0.05% Sporadic Platycrinites sp. 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Talarocrinus sp. 0.55% Sporadic 0.00% Absent 1.08% Rare 0.00% Absent 0.32% Sporadic Taxocrinus whitfieldi 1.11% Rare 0.00% Absent 0.00% Absent 0.49% Sporadic 0.58% Sporadic Tholocrinus spinosus 0.14% Sporadic 0.00% Absent 0.00% Absent 0.00% Absent 0.05% Sporadic

Table 5.3 (Continued)

Chondrichthyes Agassizodus 0.00% Absent 0.27% Sporadic 0.00% Absent 0.00% Sporadic 0.05% Sporadic Cladodus 0.00% Absent 0.27% Sporadic 0.00% Absent 0.33% Sporadic 0.16% Sporadic Ctenacanthus 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Deltodopsis 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Deltodus 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Listracanthus? 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Lophodus 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Mesodmodus 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Petrodus 0.00% Absent 0.00% Absent 0.00% Absent 9.67% Occasional 3.10% Rare Psammodus? 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Polyrhizodus 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic Taeniodus? 0.00% Absent 0.00% Absent 0.00% Absent 0.16% Sporadic 0.05% Sporadic

144 Venustodus 0.00% Absent 0.27% Sporadic 0.00% Absent 0.00% Absent 0.05% Sporadic

Grand Total: 100.00% 100.00% 100.00% 100.00% 100.00%

5.7. Statistical Estimates of Relative Abundance

In order to classify and compare faunal abundance at each quarry locality, several well-known statistical metrics were used to compare and contrast species occurrence and community composition. All statistical analyses were performed using Microsoft Excel data-analysis tools. Richness, diversity, and evenness were calculated by using census- population counts for total slab species (slab and loose samples) for both 213 and Valley

Stone quarries. In addition, the top- and bottom-of-slab communites at the 213 Quarry and the four lithofacies communities at the Valley Stone Quarry were individually analyzed for measures of richness, diversity, and evenness. All three metrics capture the range of diversity, from the number of taxa to the distribution of taxa (Patzkowsky and

Holland, 2012). In addition, the density and relative frequency were calculated for each community and for each quarry “landscape.”

5.7.1. Species Richness

Species richness is the simplest calculation in a paleoecological study

(Patzkowsky and Holland, 2012). Richness refers to the number of different species in a community or per unit area of study. However, both abundant and rare species contribute equally to the richness value. Table 5.4 shows the species richness for both the 213 and

Valley Stone quarries.

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Table 5.4 The species richness for the 213 and Valley Stone quarries and the individual communities at those localities.

Quarry, Species Location/Lithologies Richness (facies) 213 - Entire Landscape 45 213 - Top-of-slab 44 213 - Bottom-of-slab 25 Valley Stone (VS) - Entire Landscape 65 VS - Coarse-grained calcarenite 39 VS - Coarse-grained, argillaceous calcarenite 29 VS - Fine-grained calcarenites and interbedded calcilutite 21 VS - Argillaceous calcilutite 49

5.7.2. Simpson’s Index of Diversity

Simpson’s Index D (Simpson, 1949) is the probability that two random specimen samples from a community will belong to the same species. D = (∑ n*(n-1)) / N*(N-1), where n = total number of individuals of a species, and N= sum of individuals of all species. D has values ranging from 0 to 1. In contrast, Simpson’s Index of Diversity is 1 –

D, and the result will be a number between 0 and 1. The closer the number is to 1, the higher the diversity, whereas a number approaching 0 indicates lower diversity. I used an adaptation of Pearson’s “r” to interpret values between 0 and 1 (Harris et al., 2018).

Pearson’s “r”, is a statistical measurement, ranging from 0 to ±1, that provides an

146

indication of strong or weak linear relationships between two variables (Franzblau, 1962;

Hinkle et al., 1988). Table 5.5 shows the calculated Simpson’s Index of Diversity for the

213 and Valley Stone quarries.

Table 5.5 Simpson’s Index of Diversity for the 213 and Valley Stone quarries and the individual communites at those localities.

Quarry, Simpson’s Location/Lithologies Index of (facies) Diversity 213 - Entire Landscape 0.49 213 - Top-of-slab 0.63 213 - Bottom-of-slab 0.79 Valley Stone (VS) - Entire Landscape 0.90 VS - Coarse-grained calcarenite 0.89 VS - Coarse-grained, argillaceous calcarenite 0.66 VS - Fine-grained calcarenites and interbedded calcilutite 0.90 VS - Argillaceous calcilutite 0.92

5.7.3. Shannon’s Evenness Index

Shannon’s Diversity Index H was used to calculate Shannon’s Evenness Index

EH, (Shannon and Weaver, 1949), H = ∑ - (P * ln P), where P = proportion (n/N) and ln is the natural logarithm. Shannon’s Evenness Index EH equals H / ln (richness); the result will be a number between 0 and 1, and it is interpreted just like Simpson’s Index of

Diversity. A value of 0 denotes no evenness (one species is dominating the community) and 1 denotes complete evenness (the proportional abundances are nearly the same for all

147

species). In both calculations, Simpson’s Index of Diversity and Shannon’s Evenness

Index use the laws of probability and interpretations thereof; the higher the value, the more likely the outcome is to occur. In Simpson’s Index of Diversity, the total number of species (Species Richness; Table 5.4) was used in the calculations. Table 5.6 shows the calculated Shannon’s Evenness Indices for the 213 and Valley Stone quarries and their communities.

Table 5.6 Shannon’s Evenness Indices for the 213 and Valley Stone quarries and the individual communites at those localities.

Quarry, Shannon’s Location/Lithologies Evenness (facies) Index 213 - Entire Landscape 0.68 213 - Top-of-slab 0.46 213 - Bottom-of-slab 0.61 Valley Stone (VS) - Entire Landscape 0.69 VS - Coarse-grained calcarenite 0.71 VS - Coarse-grained, argillaceous calcarenite 0.57 VS - Fine-grained calcarenites and interbedded calcilutite 0.84 VS - Argillaceous calcilutite 0.76

5.7.4. Rank-Abundance Curves

Rank-abundance curves are another way to visualize the relationships between the relative abundance of each species and the number of species (Relyea and Ricklefs,

2018). The proportion was calculated for each species and then the proportions were sorted by largest to smallest to assign a rank. Rank-abundance curves are important 148

because they can easily illustrate how communities differ in species richness and species evenness.

5.7.5. Rank Abundance Curve for the 213 Quarry: Top-of-slab vs. Bottom-

of-slab Communities

In comparing curves for the fauna of the top-of-slab and bottom-of-slab communities at the 213 Quarry, the slope is very steep at the beginning of each curve

(Figure 5.51). This steepness means that the evenness for this community is very low. In other words, this community is dominated by a handful of species. This analysis allows us to compare the species richness and evenness visually on a graph. Based on the nature of the curves in Figure 5.51, the bottom-of-slab community at the 213 Quarry has a more even distribution of species, as it exhibits a lesser slope at the beginning of the curve.

213 Quarry: Top-of-slab vs. Bottom-of-slab Rank-Abundance Curve 0.7 0.6 0.5 0.4 0.3

PROPORTION 0.2 0.1 0 0 5 10 15 20 25 30 35 40 45 RANK

Top of Slab Bottom of Slab

Figure 5.51. Comparison of rank-abundance curves for fauna on top-of-slab and bottom- of-slab communities at the 213 Quarry.

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5.7.6. Rank Abundance Curve for the Valley Stone Quarry: Comparisons among the Four Lithofacies Communities

In comparing fauna of the various lithofacies communities at the Valley

Stone Quarry, the slope at the beginning of the curve for the coarse-grained,

argillaceous calcarenite community is steepest of all the communities (Figure

5.52). The steepness means that the evenness for this community is very low. This

community is dominated by a handful of species, C. subquadrata, A. leidyi and P.

bimagnaramus. Among the other three lithofacies, the fauna from argillaceous

calcilutite community is the most even (Figure 5.52).

Valley Stone Quarry Lithofacies Communities 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0

0 5 10 15 20 25 30 35 40 45 50 PROPORTION RANK

Coarse-grained calcarenite Coarse-grained, argillaceous calcarenites Fine-grained calcarenites and interbedded calcilutites Argillaceous calcilutites

Figure 5.52. Comparison of rank-abundance curves for fauna in the four lithofacies communities at the Valley Stone Quarry.

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5.7.7. Rank Abundance Curve for 213 vs. Valley Stone quarries

In comparing curves for the landscape faunas of the 213 and Valley Stone quarries, the slope is much steeper at the beginning of the curve for the 213 Quarry

(Figure 5.53), meaning that faunal evenness for this community is very low, and that the community is dominated by a handful of species, A. cf. meekanus, Rhombopora sp., and

A. leidyi. This kind of analysis allows us to compare the species richness and evenness visually on a graph, and between two locality landscapes, the fauna at the Valley Stone

Quarry is the most even (Figure 5.53).

Valley Stone vs. 213 Quarries Rank-Abundance Curve 0.60

0.50

0.40

0.30

PROPORTION 0.20

0.10

0.00 0 10 20 30 40 50 60 70 RANK

Valley Stone Quarry 213 Quarry

Figure 5.53. Comparison of rank-abundance curves for fauna from the entire landscapes of the 213 and Valley Stone quarries.

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5.7.8. Community Density

The species density formula is ∑ n / (Area), where n = total number of individuals at a given locality or community. Table 5.7 shows the calculated community density in terms of the area for each along with the number of individuals per square foot/meter for the 213 and Valley Stone quarries. The 213 Quarry collection encompassed an area of

3.25 m2 (35 ft2), whereas the Valley Stone collection encompassed an area of 4.07 m2

(43.76 ft2). Clearly, the highest density of individuals occurs in the 213 Quarry total landscape and top-of-slab communities as well as in the Valley Stone Quarry argillaceous calcilutite community. The other communities exhibit low to moderate densities.

Table 5.7 The community density for the 213 and Valley Stone quarries and the individual communities at these localities.

Community Density Quarry, Number of Number of Area Area Location/Lithologies ∑ n individuals individuals ft2 m2 (facies) per ft2 per m2 213 - Total Landscape 35 3.25 3243 92.66 997.35 213 Top-of-slab 35 3.25 2594 74.11 797.76 213 Bottom-of-slab 35 3.25 649 18.54 199.59 Valley Stone (VS) – Total Landscape 43.8 4.07 1894 43.28 465.88 VS - Coarse-grained calcarenite 21.5 1.99 723 33.67 362.47 VS - Coarse-grained, argillaceous calcarenites 9.66 0.90 376 38.92 418.97 VS - Fine-grained calcarenites and interbedded calcilutites 6.22 0.58 185 29.74 320.15 VS - Argillaceous calcilutites 6.41 0.60 610 95.16 1024.34

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5.7.9. Relative Frequency

The relative frequency formula is Relative Frequency = n/N where n = total number of individuals of a species, and N = sum of individuals of all species. The

Relative Frequency of a species is its proportion relative to the total number of all species in its community or landscape. The relative frequency represents the probability that a randomly selected sample will include that particular species. The tables for relative frequency are listed in Appendix A.

5.8. Levels of Diversity

5.8.1. Alpha diversity (α-diversity)

Alpha diversity (α-diversity) is the biodiversity within a particular area or community (Patzkowsky and Holland, 2012). The alpha diversity is simply the same as species richness (Table 5.4).

5.8.2. Beta diversity (β-diversity)

Beta diversity (β-diversity) is a measure of biodiversity made by comparing the species diversity among communities across a geographic landscape (Patzkowsky and

Holland, 2012). In other words, Beta diversity involves comparing the number of taxa that are unique to each lithofacies, community or landscape. Beta diversity is defined as the rate of change in species composition across habitats or among communities. It gives a quantitative measure of the diversity of communities and reflects change in that diversity along a gradient or landscape (Tables 5.10, 5.11 and 5.12). Tables 5.8 and 5.9 were used in defining Beta and Gamma diversity for each community and landscape.

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5.8.3. Gamma diversity (γ-diversity)

Gamma diversity refers to the total species richness over a large area or region. It is the measure of the overall diversity for all the different communities within a region. It is the product of Alpha (α diversity) of component ecosystems and the β diversity between component ecosystems. Gamma diversity can be expressed in terms of the species richness of component communities as follows: γ = S1 + S2 – c, where, S1 = the total number of species recorded in the first community, S2 = the total number of species recorded in the second community and, c = the number of species common to both communities. Table 5.8 shows the presence and absence of the species for the top-of-slab, bottom-of-slab and the entire landscape for the 213 Quarry. Table 5.9 shows the presence and absence of the species among their lithofacies communities at the Valley Stone

Quarry.

Table 5.8 Species that are present (X) vs. the species that are absent (A) for the two 213 Quarry communities.

Top-of- Bottom-of- 213 Total 213 Quarry Census Summary slab slab Landscape Anthozoa (1) Zaphrentoides spinulosus X X X

Scyphozoa? (1) Sphenothallus sp. X X X

Pelecypoda (1) Sulcatopinna missouriensis X A X

Polychaeta? (1) Crinicaminus haneyensis X A X

Ostracoda (3) 154

Table 5.8 (Continued)

Amphissites? X A X Bairdia? X A X Coronakirkbya? X A X

Articulata (6) Anthracospirifer leidyi X X X Cleiothyridina sublamellosa X X X Composita lewisensis X X X Composita subquadrata X X X Diaphragmus elegans X X X Eumetria verneuliana X A X

Inarticulata (2) Oehlertella pleurites X A X Orbiculoidea sp. X X X

Stenolaemata (9) Archimedes cf. meekanus X X X Eridopora sp. X X X Fenestella sp. X X X Glyptopora punctipora X X X Lyroporella sp. X A X Polypora sp. X X X Rhombopora sp. X X X Septopora sp. X X X Sulcoretepora sp. X X X

Blastoidea (1) Pentremites elegans X X X

Echinoidea (2) Lepidesthes formosa X A X

Archaeocidaris megastylus A X X (isolated spines only)

Crinoidea (16) Acrocrinus shumardi X A X Agassizocrinus lobatus (juvenile) X A X

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Table 5.8 (Continued)

Anartiocrinus maxvillensis X A X Aphelecrinus mundus X A X Camptocrinus cirrifer X X X Cryphiocrinus girtyi X A X Cymbiocrinus grandis X X X Cymbiocrinus tumidus X A X Dasciocrinus florealis X X X Pentaramicrinus gracilis X X X Phanocrinus bimagnaramus X X X Phanocrinus maniformis X A X Pterotocrinus acutus X X X Taxocrinus whitfieldi X X X Tholocrinus spinosus X A X Zeacrinites magnoliaeformis X A X

Chondrichthyes (2) Cladodus X A X Venustodus X A X

Species Present/Common (X): 44 25 45 Species Absent (A): 1 20 0

Table 5.9 Species that are present (X) vs. species that are absent (A) for the four lithofacies communities at the Valley Stone Quarry.

Fine-grained Coarse- calcarenites Coarse- grained, and Valley Stone Quarry Census grained argillaceous interbedded Argillaceous VS Total Summary calcarenite calcarenites calcilutites calcilutites Landscape Demospongea (1) Belemnospongia fascicularis X X X X X

Anthozoa (1) Unidentifiable tabulate coral? X A A A X

Scyphozoa? (2) Sphenothallus sp. X X X X X

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Table 5.9 (Continued)

Paraconularia missouriensis X A A X X

Pelecypoda (1) Sulcatopinna missouriensis X A A X X

Polychaeta? (1) Crinicaminus haneyensis X A A A X

Articulata (11) Anthracospirifer leidyi X X X X X Cleiothyridina sublamellosa A X X X X Composita lewisensis X X A X X Composita subquadrata X X X X X Diaphragmus elegans X X X X X Dielasma sp. A A A X X Eumetria costata X A A X X Eumetria verneuliana X X A X X Orthotetes kaskaskiensis X A A X X Punctospirifer sp. A X A X X Streptorhynchus sp. X A A A X

Inarticulata (2) Lingulipora sp. X A A A X Orbiculoidea sp. X X X A X

Stenolaemata (10) Anisotrypa sp. X A A X X Archimedes cf. meekanus X X A A X Eridopora sp. A A A X X Fenestella sp. A A X X X Fenestrellina sp. A A X A X Polypora sp. X X X X X Rhombopora sp. X X A X X Septopora sp. A A A X X Thamniscus? X X X X X Unidentifiable encrusting X X A A X bryozoan

Edrioasteroidea (2) Unidentifiable edrioasteroid X A A A X

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Table 5.9 (Continued)

Lepidodiscus laudoni A A A X X

Blastoidea (1) Pentremites elegans X X X X X

Echinoidea (1) Archaeocidaris megastylus X A X X X

Ophiuroidea (1) Unidentifiable X X X X X encrinasterid brittle star

Crinoidea (18) Agassizocrinus lobatus X X X X X Anartiocrinus maxvillensis X A X X X Aphelecrinus mundus X X A X X Camptocrinus cirrifer A X X A X Cryphiocrinus girtyi A A A X X Cymbiocrinus grandis X X A X X Cymbiocrinus tumidus X X A X X Dasciocrinus florealis X X X X X Eupachycrinus boydii A A A X X Linocrinus wachsmuthi X X X X X Pentaramicrinus gracilis X A A X X Phanocrinus bimagnaramus X X X X X Phanocrinus maniformis X A A A X Phacelocrinus longidactylus A X A A X Platycrinites sp. A A A X X Talarocrinus sp. X A X A X Taxocrinus whitfieldi X A A X X Tholocrinus spinosus X A A A X

Chondrichthyes (13) Agassizodus A X A A X Cladodus A X A X X Ctenacanthus A A A X X Deltodopsis A A A X X Deltodus A A A X X Listracanthus? A A A X X Lophodus A A A X X

158

Table 5.9 (Continued)

Mesodmodus A A A X X Petrodus A A A X X Psammodus? A A A X X Polyrhizodus A A A X X Taeniodus? A A A X X Venustodus A X A A X

Species Present/Common 39 29 21 49 65 (X): Species Absent (A): 26 36 44 16 0

The alpha, beta and gamma diversities were calculated for the 213 and Valley

Stone quarries. Tables 5.8 and 5.9 show the presence or absence of species for the 213 and Valley Stone quarries and were used to calculate the various diversities (Tables 5.10,

5.11 and 5.12). Table 5.8 shows the comparison of species presence or absence for the top-of-slab vs. the bottom-of-slab communities in the 213 Quarry, and Table 5.9 provides the same data for fauna in the four lithofacies communities at the Valley Stone Quarry.

Table 5.10 Alpha, Beta, and Gamma Diversities for the entire 213 and Valley Stone landscapes.

Alpha, Beta, and Gamma Diversity – 213 vs. Valley Stone Quarry Landscape 213 Valley Stone Community Description Quarry Quarry Diversity (species) (species)

Alpha (α) Diversity Species Richness 45 65

Change in Species Beta (β) Diversity Richness (213 vs. 44 Valley Stone)

159

Table 5.10 (Continued)

Total number of species among all lithofacies over the Gamma (γ) geographic 77 Diversity landscape (213 & Valley Stone Quarries)

Table 5.11 Alpha and Beta Diversities for top-of-slab and bottom-of-slab communities at the 213 Quarry.

Alpha, Beta, and Gamma Diversity – 213 Top-of-slab vs. Bottom-of-slab Top-of- Bottom-of - Community Diversity Description slab slab (species) (species) Alpha (α) Diversity Species Richness 44 25 Change in Species Beta (β) Diversity Richness (Top-of-slab vs. 21 Bottom-of-slab)

Table 5.12 Alpha, Beta, and Gamma Diversities for the fauna in the four lithofacies communities at the Valley Stone Quarry.

Alpha, Beta, and Gamma Diversity - VS, Coarse-grained calcarenite, coarse-grained argillaceous calcarenite, fine-grained calcarenite and interbedded calcilutite, and argillaceous calcilutite communities.

Fine- Coarse- grained Coarse- grained, calcarenites Argillaceous Community grained Description argillaceous and calcilutites Diversity calcarenite calcarenites interbedded (species) (species) (species) calcilutites (species)

Alpha (α) Species Richness 39 29 21 49 Diversity

160

Table 5.12 (Continued)

Beta (β) Change in Species Richness Between Two Communities Diversity

Coarse-grained calcarenite vs. Coarse-grained, 24 argillaceous calcarenite

Coarse-grained calcarenite vs. Fine-grained calcarenite 26 and interbedded calcilutite

Coarse-grained calcarenite 32 vs. Argillaceous calcilutite

Coarse-grained, argillaceous calcarenite vs. Fine-grained 18 calcarenite and interbedded calcilutite

Coarse-grained, argillaceous calcarenite vs. Argillaceous 35 calcilutite

Fine-grained calcarenite and interbedded calcilutite vs. 36 Argillaceous calcilutite Total number of species among all lithofacies communities over geographic landscape (Valley Stone Coarse- Gamma (γ) grained calcarenite, coarse- 65 Diversity grained argillaceous calcarenite, fine-grained calcarenite and interbedded calcilutite, and argillaceous calcilutite communities).

CHAPTER 6: DISCUSSION AND INTERPRETATIONS

Fossil echinoderm fragments are major constituents of most Paleozoic limestones; yet, complete echinoderm fossils are relatively rare in the fossil record because they are composed of many plates that disarticulate rapidly after death (e.g., Meyer, 1971b;

Baumiller, 2008). How fast echinoderms disarticulate is also related to the amount of exposure prior to burial (Brett et al., 1997; Kidwell, 2013). Hence, when we find localities with relatively abundant echinoderms, like the two in this study, it is important for future work to understand the conditions that permitted so many echinoderms to live in one place, as well as the conditions that permitted their largely intact preservation.

6.1. Regional Geologic Controls

During Late Mississippian time, the studied localities in the Ramey Creek

Member and their fossils were parts of seafloor communities that developed in a shallow, open-marine setting on south-central parts of Laurussia (Figure 2.5 and 2.6). The communities were located at about 20° south latitude in an arid, subtropical belt (Boucot et al., 2013) that was favorable for the deposition of carbonate sediments and for organisms that secreted calcareous skeletons (Lees, 1975). Although the Ramey Creek

Member was part of one of the many local transgressive cycles (Figure 2.1), global sea level was decreasing at the time due to the inception of southern-hemisphere Gondwana glaciation (Haq and Schutter, 2008) (Figure 2.5).

The structural/tectonic framework was also instrumental in generating the regional depositional setting. The Waverly Arch and subsurface portions of the Kentucky

River Fault Zone, the northern bounding fault of the Rome Trough (Figure 2.3), reflect

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basement structures that crossed the study area between the two studied sections.

Previous work by Ettensohn (1975b, 1977, 1980, 1981b, 1986), Dever (1980, 1990),

Dever et al. (1977, 1990), and Ettensohn and Dever (1979) has shown that these structures were periodically reactivated during Mississippian time, and that the northern area was continually uplifted relative to areas south of the Kentucky River Fault Zone. In support of these interpretations, these workers highlighted the presence of disjunct unit distribution, thinning units, abrupt facies changes near structures, unexpected absence of units, deep erosional truncation, and prominent disconformities. Ettensohn (1975b) also pointed out that the Ramey Creek Member facies north of the fault zone contained more coarse-grained calcarenites and probably reflected higher-energy environments than those that occur south of the fault zone, and I have made the same observation relative to the two studied localities (Figure 2.4). The Valley Stone Quarry locality occurs north of the fault zone and shows more coarse-grained, calcarenitic, shoal facies (Figure 6.1) than does the 213 Quarry locality to the south.

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Figure 6.1. Schematic representation of upper Slade Formation transgressive environments relative to regional structures in east-central Kentucky. The approximate locations of the 213 and Valley Stone quarries are shown by red stars (modified after Ettensohn, 1975b, 1980).

6.2. Lithofacies and Depositional Environments

6.2.1. Valley Stone Quarry Lithofacies

Ettensohn (1975b, 1977) indicated that all of the Slade members north of the

Kentucky River Fault Zone had more shallow-water features and facies than those to the south. The Ramey Creek Member represents shallow, open-marine, outer-ramp deposition just seaward of the Tygarts Creek sandbelt (Figures 3.21 and 6.1), and this situation is supported by its stratigraphic position above the Tygarts Creek (Figures 3.2 and 6.2) but below the Maddox Branch in an upwardly transgressing sequence (Figures

2.1 and 3.21). However, in contrast to the situation at the 213 Quarry, Ettensohn (1975a) also showed that the Ramey Creek Member on the northern block was largely composed of an irregular array of tidally and storm-wave-influenced shoals, where coarse skeletal sands were deposited, and deeper intervening basins where calcareous mud and silt, as well as argillaceous sands, predominated (Figure 6.3). The coarse skeletal sands were deposited at or near normal wave base, whereas argillaceous calcarenites, calcisiltites, and calcilutites with interbedded shales and mudstones were deposited below tidal and wave influence. Storm waves, however, may have periodically penetrated much deeper, destroying fossil communities and mixing the muds and skeletal sands. A probably analogous array of shoals and basins with intervening facies is present in shallow-marine environments, 60–100-m deep, in the Persian Gulf (Kassler, 1973). As in the Persian

Gulf (Kassler, 1973), the interaction of waves and tides with organism-produced sediment and regional structures probably generated the array of shoals, basins and intervening facies (Figure 6.3) that have thus far been referred to as Ramey Creek

Member lithofacies, based on lithology in previous parts of the study. In the following

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four sections, these lithofacies will be interpreted as depositional environments with faunal associations.

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Figure 6.2. Four distinct lithofacies and their interpreted environments for the Valley Stone Quarry (Figures 1.6 and 3.4). Fine dashed lines in the shoal lithofacies represent scours and dunes.

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Figure 6.3. Interpretive drawing of the four lithofacies and their depositional settings in the Ramey Creek Member at the Valley Stone Quarry. Red line across the top represents sea level, whereas the blue line represents normal wave base. The darker teal color at the bottom and left represents the Tygarts Creek Member. The entire diagram suggests facies migration to the left with rising sea level.

6.2.1.1. Shoal Lithofacies

Although previous interpretations of the Ramey Creek Member indicate that it represents environments below normal wave base (Figure 3.20 and 3.21; Ettensohn,

1975b, 1980, 1981a, b), storms apparently reworked these sediments frequently, especially those near normal wave base (e.g., Aigner, 1982; Marsaglia and Klein, 1983;

Duke, 1985; Finnegan and Droser, 2008) as shown by the nature of some of the sediments and sedimentary structures, like crossbeds, crude graded bedding, scours, rip- up clasts, and ripple marks (Figures 3.5 and 6.4). Throughout the Ramey Creek Member, especially on the uplifted northern block, beds of clean, well-washed skeletal sands are a common lithofacies (Figures 2.4, 3.2, and 3.7). The lack of any mud in these sands suggest that they were deposited in a high-energy environment where continual winnowing washed any muds into deeper environments. In addition, the presence of dune-like bedforms and basin-like scours (Figures 3.4 and 6.2) suggests that the sands were continually reworked into migrating shoals. When complete fossil assemblages are preserved in this lithology, like the delicate echinoderms in this study (Figures 6.45 and

6.6), the detailed preservation suggests rapid burial by migrating sands. Bioturbation is typically lacking in this facies because the sands are too mobile for organisms to establish themselves. Hence, this lithofacies (coarse-grained calcarenite lithofacies) represents a lacework of migrating sand shoals at or near normal wave base as previously interpreted or slightly shallower that previously reported (Figure 6.3; Table 6.1).

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Figure 6.4. A close-up view of the shoal and shoal-margin depositional environments in the Ramey Creek Member. Fine dotted lines in shoal represent scours and cross-bedded dunes. Note shale beds and partings in the shoal-margin environment.

Table 6.1 Environmental Interpretations for lithofacies at 213 and Valley Stone quarries.

213 & Valley Stone Quarries Environmental Interpretations Environment by Environmental Lithology/Lithofacies Interpretation 213 Quarry - Thin-bedded, Storm-influenced argillaceous, fine-grained ramp; comparable calcarenite and interbedded to Transitional gray-green shale below. Valley Stone Quarry - Coarse-grained, calcarenite Shoal Valley Stone Quarry - Coarse-grained, argillaceous calcarenite Shoal-Margin

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Table 6.1 (Continued) Valley Stone Quarry - Fine- grained calcarenite, and interbedded calcilutite Transitional Valley Stone Quarry - Argillaceous calcilutite Basinal

Figure 6.5. Randomly oriented fossil fragments (crinoids, brachiopods, sponges, and blastoid) from the weathered surface of a stylolitic bedding plane from the coarse-grained calcarenite lithofacies (shoal environment) at the Valley Stone Quarry. The crinoid calyx in the upper right is complete but not attached to its stem.

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Figure 6.6. A well-preserved crinoid, Aphelocrinus mundus, on a weathered stylolite surface from coarse-grained calcarenite lithofacies, or shoal environment, in the Valley Stone Quarry.

6.2.1.2. Shoal-margin Lithofacies

The coarse-grained, argillaceous calcarenite lithofacies differs from the coarse- grained calcarenite lithofacies in the presence of an argillaceous and calcareous muddy matrix within the calcarenites and the presence of many shale interbeds (Figure 6.4). The facies is typically more even-bedded and lacks the many scours and mound-like bedforms found in the coarse-grained calcarenites (see Figure 6.4). Moreover, in this lithofacies, the fossils are typically more complete and better preserved in the shaly upper parts of the beds, and bioturbation is relatively common. The coarse-grained sands in this lithofacies apparently represent storm-backflow deposits (Aigner, 1982) derived from adjacent

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shoals, and because they were deposited further below wave base, muddy matrix materials are more common. Because the effects of storms were less frequent further below wave base, organism communities could become established on the firm substrates, but periodically mud fallout from nearby storms would rapidly smother and bury these communities. Because of stratigraphic position above the shoal facies and the presence of muds as matrix and interbeds, I interpret this facies to represent shoal-margin environments that were deeper and more distal to the shoals (Figures 6.2 and 6.3; Table

6.1).

6.2.1.3. Transitional Lithofacies

In this lithofacies, the rocks are composed of a fine-grained calcarenites with interbedded calcisiltites/calcilutite and shales (Figure 6.2); bioturbation may be intense.

High-energy bedforms are rare, and organisms more abundant here, presumably because of deeper, more stable conditions. In this setting only the finer-grained, most distal parts of the storm backflow deposits were transported into the environment. Based on stratigraphic position and the nature of sedimentation, this environment is interpreted as transitional into the deepest-water or basinal environments (Figures 6.2 and 6.3; Table

6.1).

6.2.1.4. Basinal Lithofacies

Sediments in the argillaceous calcilutite and shale lithofacies (Figure 6.2) are finer-grained than any other sediments in the Ramey Creek Member. In places, the calcilutites are nodular and interbedded with shales. Fossils are more abundant than in any other lithofacies, and many were specially adapted to muddy substrates. Bioturbation

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may be intense and is probably responsible for the nodular nature of many calcilutites.

This lithofacies was probably deeper and more distal from the shoals than any of the other lithofacies, which accounts for its predominantly fine-grained nature. The environmental stability accompanying its distal nature and greater depth also probably accounts for the abundance of fossils and bioturbation. Based on its stratigraphic position and the features noted above, this lithofacies is interpreted to represent a deeper-water basinal environment (Figures 6.2 and 6.3; Table 6.1).

6.2.2. The 213 Quarry Lithofacies

Previous research by Ettensohn (1975b, 1977, 1980, 1981b, 1986), Dever (1980,

1990), Dever et al. (1977, 1990), and Ettensohn and Dever (1979) has inferred changes across the Kentucky River Fault Zone such that 213 Quarry lithologies probably developed on the down-dropped block (Figure 6.1), where conditions were generally deeper and less energetic. In contrast to the northern platform-like area where a shallow open-marine facies mosaic developed (Valley Stone Quarry; Figures 6.1 and 6.3), the 213

Quarry locality apparently developed on a gentle eastward-dipping ramp (Figure 6.1). A single lithofacies, represented by thin-bedded, fine-grained, argillaceous calcarenites and interbedded shales (Figures 3.13, 3.14 and 3.18) characterizes this locality. It is comparable to the fine-grained calcarenite, and interbedded calcilutite lithofacies in the

Valley Stone Quarry (Table 6.1).

The individual layers in this 213 Quarry lithofacies range from 0.5–7.5-cm thick, and often exhibit graded bedding, cross bedding, scours and gutter casts (Figures 3.14,

3.17 and 3.18), indicating that they were storm-related deposits or tempestites (e.g.,

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Aigner, 1982). As storms hit shallower environments, they would have generated high- energy backflow currents that transported sands basinward down the ramp. In areas that had not experienced storms and backflow currents for a while, firm substrates capable of supporting organism communities would have developed, and on these substrates small patchy bryozoan-brachiopod-crinoid filter-feeding communities, like the one studied here, apparently developed. The area of the studied community, approximately 3.3 m2 (35 ft2) gives an idea of about how large these communities would have been. Because this was a ramp subject to frequent storm activity, communities like this would have periodically been buried below the backflow deposits if they were proximal to the storm.

On the other hand, if they were more distal to storms, they may have been buried by clouds of mud wafting away from storms occurring elsewhere on the ramp.

At the 213 Quarry, a distinct history of depositional events can be established based on the sedimentary evidence from the calcarenite layer on which the fossils were found (Harris and Ettensohn, 2015a): 1.) On the bases of many of the 213 slabs, a prominent basal deposit of breccia and rip-up clasts occurs along with scours, slickensides, brittle fractures (Figures 3.13, and 6.7–6.11). These features have been interpreted to represent a locally catastrophic event, such as a storm or seismic event. 2.)

A firm substrate developed on this material, and the bottom-of-slab community developed. 3.) The bottom-of-slab community was subsequently knocked down and transported a short distance. No crinoid holdfasts were found, so the crinoids were apparently ripped from the bottoms with short stem segments. The Archimedes bryozoans also lack holdfasts as well as attached fronds, so they were also transported, but not far enough for abrasion to have occurred. The state of these organisms suggests that they

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were knocked down by powerful currents from a nearby storm and buried by the attendant mud fallout (Harris and Ettensohn, 2015b). 4.) The bottom-of-slab community, now encased in mud, was later buried by another storm deposit or tempestite, indicated by crossbedding, graded bedding, and scours (Figure 3.18). 5.) A firm substrate later developed on this tempestite, and a second larger bryozoan-brachiopod-crinoid community developed on top of it (Figure 3.15). 6.) The second community was knocked down by high-energy currents from a nearby storm event based on the same criteria noted above in the third event. 7.) Based on the disarticulation of some of the crinoids (Figure

6.12) this community was exposed briefly, perhaps for up to a week based on the work of

Meyer (1971b). 8.) The second, or top-of-slab community, was then buried by mud fallout from an adjacent storm (Figure 1.7), similar to the situation noted in the fourth event above. The result of these eight events is a series of slabs with bottom-of-slab and top-of-slab communities (Figure 3.18). The bottom of slab community is effectively

“welded” to the bottom of the bed. Although I have treated the two communities as separate, based on the community compositions, it is possible they may be related. Figure

6.13 shows the two possibilities. Figure 6.13A shows the communities as individual communities separated by a major tempestite layer, whereas Figure 6.13B shows that only part of the bottom-of-slab community was knocked down and buried by the tempestites layer, allowing unburied parts to regenerate and expand onto the top of the tempestite layer, thereby forming the second or top-of-slab community. Some of the slabs showing the top-of-slab community are thin enough (Figure 3.14) that this could have been a possibility, but firm evidence is lacking.

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Figure 6.7. Brittle fractures and “slickensides” from the base of a slab from the 213 Quarry.

Figure 6.8. Rip-up clasts on the bottom-of-slab from the 213 Quarry.

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Figure 6.9. An example of rip-clasts and load structures on the base of a slab from the 213 Quarry.

Figure 6.10. An example of rip-up clasts, scoured base and storm layer from the 213 Quarry.

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Figure 6.11. An example of ripped up crinoids, scour (red line) and disorganized sediment on the base of a slab from the 213 Quarry.

Figure 6.12. The flexible crinoid, Taxocrinus whitfieldi, on two slabs (separated below scale). Calyx on the left shows disarticulation of the calyx and stem. Calyx on the right shows a largely intact calyx and stem. Single valve of Anthracospirifer leidyi on lower right.

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Figure 6.13. Two possibilities for relationships between the top- and bottom-of-slab communities at the 213 Quarry: A.) Top- and bottom-of slab communities (mixed fossils) are unrelated, having been separated in time and space by a graded tempestite (beds with decreasing circle sizes); B.) Top- and bottom-of-slab communities are initially separated by a tempestite bed that terminates to the left such that the communities merge into one.

6.3. Taphonomy

The effects of preservational mode, or how an organism entered the fossil record, must be taken into account before we can interpret how an organism lived and what its community was like. In particular, taphonomy is concerned with the way in which dead organisms enter the fossil record, and deals with all the post-mortem events experienced by organisms, including biostratinomy and diagenesis. In particular, biostratinomy is the part of taphonomy that deals with events after death and before final burial, and much of this section deals with biostratinomic interpretations for communities at the 213 and

Valley Stone quarries. The most valuable fossil communities are those that are overwhelmed by catastrophes that “freeze” the assemblage on a bedding plane. Such

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assemblages are called census assemblages or life assemblages, which is what appears to be present at the quarry localities. However, are these really life assemblages, which were buried in place, and what is the evidence for one interpretation or the other?

6.3.1. Biostratinomic Indicators for the 213 Quarry communities

The indicators discussed herein will deal with both the bottom-of-slab and top-of- slab communities and include faunal composition, fossil density, size-frequency distribution, dissociation of hard parts, surface conditions of fossils, orientation of fossils, and the chemical and mineral conditions of the hardparts (e.g., Ager, 1963; Dodd and

Stanton, 1981). With faunal composition, we ask the question as to whether all of the life modes and tolerances of the community organisms are compatible. In the case of the 213

Quarry communities, most of the organisms were filter feeders (Table 6.2), which could have partitioned the water column by tiering, so that competition was not a major problem (Tables 6.2, 6.3 and 6.4; Figure 6.14).

Table 6.2 Summary of the top three most abundant species within each community.

Total Quarry, Top three most abundant species (Relative Percentage Location/Lithologies Frequency) of (facies) Community Archimedes cf. Anthracospirifer 213 Quarry - Entire Rhombopora sp. meekanus leidyi 76.65% Landscape (13.32%) (53.59%) (9.74%) Archimedes cf. Anthracospirifer 213 Quarry - Top-of- Rhombopora sp. meekanus leidyi 76.60% slab (10.06%) (58.79%) (7.75%)

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Table 6.2 (Continued)

Archimedes cf. Anthracospirifer 213 Quarry - Bottom- Rhombopora sp. meekanus leidyi 76.89% of-slab (26.35%) (32.82%) (17.72%) Valley Stone (VS) Composita Anthracospirifer Belemnospongia Quarry - Entire subquadrata leidyi fascicularis 48.18% Landscape (21.59%) (14.32%) (12.27%) Anthracospirifer Belemnospongia Phanocrinus VS - Coarse-grained leidyi fascicularis bimagnaramus 50.83% calcarenite (Shoal) (22.93%) (18.51%) (9.39%) VS - Coarse-grained, Composita Anthracospirifer Phanocrinus argillaceous subquadrata leidyi bimagnaramus 70.47% calcarenites (Shoal- (57.18%) (6.91%) (6.38%) Margin VS - Fine-grained calcarenites and Anthracospirifer Phanocrinus Composita interbedded leidyi bimagnaramus subquadrata 47.10% calcilutites (20.11%) (13.76%) (13.23%) (Transitional) Composita Belemnospongia VS - Argillaceous Pentremites subquadrata fascicularis 38.04% calcilutites (Basinal) elegans (8.20%) (17.54%) (12.30%)

Table 6.3 Interpreted feeding modes for the 213 Quarry.

Interpreted Feeding Modes for the 213 Quarry Constituent Phylum Class 213 Quarry Genera/Species Porifera Demospongea Belemnospongia fascicularis Absent Unidentifiable tabulate Anthozoa coral? Absent Zaphrentoides spinulosus Passive Predator Cnidaria Passive Sphenothallus sp. Predator/Filter Scyphozoa? Feeder? Paraconularia missouriensis Absent

Mollusca Pelecypoda Sulcatopinna missouriensis Filter Feeder

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Table 6.3 (Continued)

Annelida? Polychaeta? Crinicaminus haneyensis Passive Predator/ Deposit Browser/Deposit/ Amphissites? Scavenger Browser/Deposit/ Arthropoda Ostracoda Bairdia? Scavenger Browser/Deposit/ Coronakirkbya? Scavenger Anthracospirifer leidyi Filter Feeder Cleiothyridina sublamellosa Filter Feeder Composita lewisensis Filter Feeder Composita subquadrata Filter Feeder Diaphragmus elegans Filter Feeder Articulata Dielasma sp. Absent Eumetria costata Absent Brachiopoda Eumetria verneuliana Filter Feeder Orthotetes kaskaskiensis Absent Punctospirifer sp. Absent Streptorhynchus sp. Absent Lingulipora sp. Absent Inarticulata Oehlertella pleurites Filter Feeder Orbiculoidea sp. Filter Feeder Anisotrypa sp. Absent Archimedes cf. meekanus Filter Feeder Eridopora sp. Filter Feeder Fenestella sp. Filter Feeder Fenestrellina sp. Absent Glyptopora punctipora Filter Feeder Lyroporella sp. Bryozoa Stenolaemata Filter Feeder Polypora sp. Filter Feeder Rhombopora sp. Filter Feeder Septopora sp. Filter Feeder Sulcoretepora sp. Filter Feeder Thamniscus? Filter Feeder Unidentifiable encrusting bryozoan Absent

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Table 6.3 (Continued)

Unidentifiable edrioasteroid Absent Edrioasteroidea Lepidodiscus laudoni Absent Blastoidea Pentremities elegans Filter Feeder Lepidesthes formosa Scavenger/Deposit Echinoidea Archaeocidaris megastylus Scavenger/Deposit Ophiuroidea Unidentifiable encrinasterid Absent Acrocrinus shumardi Filter Feeder Agassizocrinus lobatus Filter Feeder Anartiocrinus maxvillensis Filter Feeder Aphelecrinus mundus Filter Feeder Camptocrinus cirrifer Filter Feeder Cryphiocrinus girtyi Filter Feeder Cymbiocrinus grandis Filter Feeder Cymbiocrinus tumidus Filter Feeder Echinodermata Dasciocrinus florealis Filter Feeder Eupachycrinus boydii Absent Linocrinus wachsmuthi Absent Crinoidea Pentaramicrinus gracilis Filter Feeder Phacelocrinus longidactylus Absent Phanocrinus bimagnaramus Filter Feeder Phanocrinus maniformis Filter Feeder Platycrinites sp. Absent Pterotocrinus acutus Filter Feeder Talarocrinus sp. Absent Filter Taxocrinus whitfieldi Feeder/Passive Predator Tholocrinus spinosus Filter Feeder Zeacrinites magnoliaeformis Filter Feeder

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Table 6.3 (Continued)

Agassizodus Absent Cladodus Predator Ctenacanthus Absent Deltodopsis Absent Deltodus Absent Chondrichthyes Listracanthus? Absent (teeth) Lophodus Absent Chordata Mesodmodus Absent Psammodus? Absent Polyrhizodus Absent Taeniodus? Absent Venustodus Predator Chondrichthyes (dermal skin Petrodus Absent plates)

Table 6.4 Interpreted feeding tiers for the 213 Quarry.

Interpreted Feeding Tiers for the 213 Quarry Constituent Phylum Class 213 Quarry Genera/Species Porifera Demospongea Belemnospongia fascicularis Absent Unidentifiable tabulate Anthozoa coral? Absent Zaphrentoides spinulosus Lower Cnidaria Sphenothallus sp. Lower/ Intermediate Scyphozoa? Paraconularia missouriensis Absent Mollusca Pelecypoda Sulcatopinna missouriensis Intermediate Annelida? Polychaeta? Crinicaminus haneyensis Lower Amphissites? Epiphyte/ Epizoan Arthropoda Ostracoda Bairdia? Epiphyte/ Epizoan Coronakirkbya? Epiphyte/ Epizoan

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Table 6.4 (Continued)

Anthracospirifer leidyi Lower Cleiothyridina sublamellosa Lower Composita lewisensis Lower Composita subquadrata Lower Diaphragmus elegans Lower Articulata Dielasma sp. Absent Eumetria costata Absent Brachiopoda Eumetria verneuliana Lower Orthotetes kaskaskiensis Absent Punctospirifer sp. Absent Streptorhynchus sp. Absent Lingulipora sp. Absent Inarticulata Oehlertella pleurites Lower Orbiculoidea sp. Lower Anisotrypa sp. Absent Archimedes cf. meekanus Intermediate Eridopora sp. Lower Fenestella sp. Lower Fenestrellina sp. Absent Glyptopora punctipora Lower Lyroporella sp. Lower Bryozoa Stenolaemata Polypora sp. Lower Rhombopora sp. Lower Septopora sp. Lower Sulcoretepora sp. Lower Thamniscus? Absent Unidentifiable encrusting bryozoan Absent

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Table 6.4 (Continued)

Unidentifiable edrioasteroid Absent Edrioasteroidea Lepidodiscus laudoni Absent Blastoidea Pentremities elegans Intermediate Lepidesthes formosa Lower Echinoidea Archaeocidaris megastylus Lower Ophiuroidea Unidentifiable encrinasterid Absent Acrocrinus shumardi Upper Agassizocrinus lobatus Lower Anartiocrinus maxvillensis Upper Aphelecrinus mundus Upper Camptocrinus cirrifer Intermediate Cryphiocrinus girtyi Intermediate Cymbiocrinus grandis Intermediate Echinodermata Cymbiocrinus tumidus Intermediate Dasciocrinus florealis Intermediate Eupachycrinus boydii Absent Crinoidea Linocrinus wachsmuthi Absent Pentaramicrinus gracilis Intermediate Phacelocrinus longidactylus Absent Phanocrinus bimagnaramus Intermediate Phanocrinus maniformis Intermediate Platycrinites sp. Absent Pterotocrinus acutus Lower Talarocrinus sp. Absent Taxocrinus whitfieldi Intermediate Tholocrinus spinosus Intermediate Zeacrinites magnoliaeformis Intermediate

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Table 6.4 (Continued)

Agassizodus Absent Cladodus Nektic Ctenacanthus Absent Deltodopsis Absent Deltodus Absent Chondrichthyes Listracanthus? Absent (teeth) Lophodus Absent Chordata Mesodmodus Absent Psammodus? Absent Polyrhizodus Absent Taeniodus? Absent Venustodus Nektic Chondrichthyes (dermal skin Petrodus Absent plates)

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Figure 6.14. Schematic reconstruction of the before-burial top-of-slab community in the Ramey Creek Member at the 213 Quarry. Bottom of slab-community would be preserved just below the shale break. Organism rank and abundance in the figure are based on species frequencies in Table 5.2.

A few passive predators, browsers/scavengers were also present (Table 6.2), but based on similar Chesterian crinoid/bryozoan communities (e.g., Chesnut and Ettensohn,

1988), none of the organisms or their tolerances were incompatible. In fact, level-bottom communities like those in the 213 Quarry are typically dominated by detritus-feeding organisms, filter and deposit feeders, and commonly aggregate together in clusters

(Speden, 1966). All of the 213 communities show high organism densities (Table 5.7), but these high densities are almost certainly not related to slow sedimentation over very long periods of time. In contrast, the 213 communities developed on firm tempestite surfaces that were deposited during major hurricane-type storms (e.g., Marsaglia and

Klein, 1983). If these storms exhibited the same types of periodicity that we currently see in hurricanes, then storms may have crossed the same areas every few years to a few tens of years, suggesting a time frame across which the 213 communities developed. Hence, it seems likely that the 213 quarry communities developed their relatively high standing densities through organism clustering at one place over a few tens of years, contradicting the possibility of major time averaging during long periods. Each 213 community is represented by one thin layer of compatible organisms that appear to have developed over a few tens of years between major storm events. No evidence exists for the presence of organisms transported in from other communities, and the time frame is too short for major time averaging of organisms. The only possible incompatible fossils present in the community were a few chondrichthyan teeth, which were apparently washed in, as the sharks were never parts of these benthic communities.

If the 213 communities were once life assemblages or census assemblages, typical size-frequency distributions suggest that all life stages should be represented in the

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communities. However, because of size constraints on preservation potential during transportation and decreased longevity for gerontic individuals, the typical size-frequency for invertebrate communities should show lower numbers of juvenile and gerontic individuals and higher numbers of average-size adults (Ager, 1963; Fagerstrom, 1964).

Using size as a proxy for life stage, the size-frequency graphs for two of the community species, Taxocrinus whitfieldi and Pentremites elegans, show just such patterns (Figures

6.15 and 6.16) , indicating that they were probably derived from a single community or a single event.

The conditions of organism hard parts from the communities is another useful biostratinomic indicator. Dissociation of shells, in particular, suggests transportation, but nearly all brachiopod shells in the 213 communities were fully articulated and smashed, suggesting that they were buried while alive and were buried fast enough that sediment could not infill the valves, thus contributing to their smashed state as a result of later compaction. Skeletal fragmentation is another critical factor, but the only seriously fragmented fossils in the communities were the bryozoans, especially Archimedes cf. meekanus. Although the central calcareous spire is everywhere preserved (Figures 1.7 and 3.15), the more delicate trellis-like fronds are typically broken and shredded into fine debris. In addition, the root-like holdfasts of Archimedes are always broken and shattered; none were found in place. The Archimedes spires, moreover, which have a prominent linear aspect, are typically randomly oriented, but in a few places show two directions of preferential orientation (Figure 3.15), suggesting transportation.

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213 Quarry: Histogram of Taxocrinus whitfieldi Cup Width (cm) Normal

18 Mean 1.621 StDev 0.4233 16 N 33

n 10

e 8

F 6

0 1.0 1.5 2.0 2.5 Cup Width (cm)

Figure 6.15. Size-frequency histogram of the flexible crinoid, Taxocrinus whitfieldi, cup width for the 213 Quarry communities. The red curve shows how a normal distribution for the collection of specimens would appear.

213 Quarry: Histogram of Pentremites elegans Theca Height (cm) Normal 25 Mean 1.309 StDev 0.5775 N 34 20

c 15

F 10

0 0 1 2 3 Theca Height (cm)

Figure 6.16. Size-frequency histogram of the blastoid, Pentremites elegans, theca height from the two communities in the 213 Quarry. The red curve shows how a normal distribution for the collection of specimens would appear.

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Likewise, although most of the crinoids and blastoids show nearly perfect preservation of crowns and thecae with pinnules and brachioles intact (Figure 3.15), their stems were always broken (Figure 3.15), and very few holdfasts were found; no attached crinoids or blastoids were found. This state of preservation suggests that the crinoids and blastoids were ripped up and transported, although the preservation of delicate parts suggests that they were not transported far. Most of the crinoids calyces were found intact, although a few Taxocrcinus calyces show various states of in-place disarticulation

(Figure 6.12), suggesting that they lay exposed on the surface for some time before final burial. Based on disarticulation studies by Meyer (1971a, b), which show modern crinoid disarticulation within 5-6 days, at least some of the crinoids must have laid exposed on the sea bottom after transportation for 5-6 days before final burial. In addition, none of the fossils in these communities show any signs of corrosion, abrasion or encrustation by organisms, suggesting that burial, although not immediate, was relatively rapid. As the organisms were apparently buried relatively fast, any chemical alteration of the hardparts is not a major consideration. The only indication of chemical alteration occurs with the

Sulcatopinna shells, which are largely dissolved and preserved as external molds. This is a common and expected alteration, as Sulcatopinna was a pelecypod with aragonitic shells, and aragonite quickly dissolves or is replaced after an organism dies.

The above biostratinomic indicators suggest that the 213 Quarry fossil organisms were likely derived from the same community, and that they were ripped up from their original substrate and transported a short distance by currents associated with a nearby storm. Unattached or loosely attached organisms like the brachiopods, corals, sphenothallids and Sulcatopinna were merely picked up and moved, whereas the more

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firmly attached organisms, like crinoids, blastoids and bryozoans, were ripped from their holdfasts and moved along with the others. Any delicate, brittle structures, like the

Archimedes fronds, were apparently highly fragmented during transportation, whereas those delicate structures sheathed in connective tissue, like the plates, pinnules and brachioles of crinoids and blastoids were transported intact. Very small or soft-bodied organisms were probably picked up and transported to areas farther away as suspended materials. At least some of the crinoids show evidence of in-place disarticulation, indicating that they may have laid exposed for several days on the sea bottom, before being buried by a suspended mud fallout, accompanying a more distal storm event.

Sometime later, perhaps during the same or a later storm season, the mud-covered organisms were buried below a proximal tempestite, or storm back-flow deposit, accompanying a nearby storm. Hence, the 213 Quarry assemblages do probably approximate the makeup of the original community. Using the terminology of Fagerstrom

(1964), the 213 Quarry organisms represent “transported fossil assemblages,” and they appear to have been contemporaneous, from the same community, and the product of mass mortality. Study of such death assemblages, even though transported, suggests that they still provide strong signals of original community composition and environmental dynamics, as well as species rank and abundance (Kidwell and Flessa, 1996; Kidwell,

2001). Based on the relative abundance of species/genera in the 213 Quarry communities

(Tables 4.2, 4.3, 4.12 and 5.2), the original, before-transportation, 213 community may have appeared as reconstructed in Figure 6.14.

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6.3.2. Biostratinomic Indicators for the Valley Stone Quarry communities

In contrast to the 213 Quarry, where all specimens were collected from one of two bedding planes (bottom- and top-of-slab communities), specimens from the Valley Stone

Quarry were collected from slabs that were generated randomly during quarry blasting.

Only later were the slabs and their contained fossils characterized by lithofacies. This means that the exact location or orientation of the specimens within the quarry are uncertain, and that they are certainly coming from multiple bedding planes in any one lithofacies. As a consequence, we cannot be as certain about taphonomic indicators from the Valley Stone Quarry, and any interpretations are generalized by lithofacies.

Nonetheless, similar to the situation at the 213 Quarry, most of the organisms in all environments were filter feeders (Table 6.2), which presumably partitioned the water column by tiering, so that competition was not a major problem (Tables 6.2, 6.5 and 6.6;

Figure 6.20).

Table 6.5 Shows the interpreted feeding modes for the Valley Stone Quarry.

Interpreted Feeding Modes for the Valley Stone Quarry Constituent Phylum Class Valley Stone Genera/Species Quarry Porifera Demospongea Belemnospongia fascicularis Filter Feeder Unidentifiable tabulate Anthozoa coral? Filter Feeder Zaphrentoides spinulosus Absent Passive Cnidaria Sphenothallus sp. Predator/Filter Scyphozoa? Feeder? Paraconularia missouriensis Planktic Predator Mollusca Pelecypoda Sulcatopinna missouriensis Filter Feeder

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Table 6.5 (Continued)

Passive Crinicaminus haneyensis Predator/ Annelida? Polychaeta? Deposit Feeder Amphissites? Absent Arthropoda Ostracoda Bairdia? Absent Coronakirkbya? Absent Anthracospirifer leidyi Filter Feeder Cleiothyridina sublamellosa Filter Feeder Composita lewisensis Filter Feeder Composita subquadrata Filter Feeder Diaphragmus elegans Filter Feeder Articulata Dielasma sp. Filter Feeder Eumetria costata Filter Feeder Brachiopoda Eumetria verneuliana Filter Feeder Orthotetes kaskaskiensis Filter Feeder Punctospirifer sp. Filter Feeder Streptorhynchus sp. Filter Feeder Lingulipora sp. Filter Feeder Inarticulata Oehlertella pleurites Absent Orbiculoidea sp. Filter Feeder Anisotrypa sp. Filter Feeder Archimedes cf. meekanus Filter Feeder Eridopora sp. Filter Feeder Fenestella sp. Filter Feeder Fenestrellina sp. Filter Feeder Glyptopora punctipora Absent Lyroporella sp. Bryozoa Stenolaemata Absent Polypora sp. Filter Feeder Rhombopora sp. Filter Feeder Septopora sp. Filter Feeder Sulcoretepora sp. Absent Thamniscus? Filter Feeder Unidentifiable encrusting bryozoan Filter Feeder

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Table 6.5 (Continued)

Unidentifiable edrioasteroid Filter Feeder Edrioasteroidea Lepidodiscus laudoni Filter Feeder Blastoidea Pentremities elegans Filter Feeder Lepidesthes formosa Absent Echinoidea Scavenger/ Archaeocidaris megastylus Deposit Scavenger/ Unidentifiable encrinasterid Predator/Deposit Ophiuroidea Feeder Acrocrinus shumardi Absent Agassizocrinus lobatus Filter Feeder Anartiocrinus maxvillensis Filter Feeder Aphelecrinus mundus Filter Feeder Camptocrinus cirrifer Filter Feeder Cryphiocrinus girtyi Filter Feeder Cymbiocrinus grandis Filter Feeder Echinodermata Cymbiocrinus tumidus Filter Feeder Dasciocrinus florealis Filter Feeder Eupachycrinus boydii Filter Feeder Linocrinus wachsmuthi Filter Feeder Crinoidea Pentaramicrinus gracilis Filter Feeder Phacelocrinus longidactylus Filter Feeder Phanocrinus bimagnaramus Filter Feeder Phanocrinus maniformis Filter Feeder Platycrinites sp. Filter Feeder Pterotocrinus acutus Absent Talarocrinus sp. Filter Feeder Filter Taxocrinus whitfieldi Feeder/Passive Predator Tholocrinus spinosus Filter Feeder Zeacrinites magnoliaeformis Absent

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Table 6.5 (Continued)

Agassizodus Predator Cladodus Predator Ctenacanthus Predator Deltodopsis Predator Deltodus Predator Chondrichthyes Listracanthus? Predator (teeth) Lophodus Predator Chordata Mesodmodus Predator Psammodus? Predator Polyrhizodus Predator Taeniodus? Predator Venustodus Predator Chondrichthyes (dermal skin Petrodus Predator plates)

Table 6.6 Interpreted feeding tiers for the Valley Stone Quarry.

Interpreted Feeding Tiers for the Valley Stone Quarry Valley Phylum Class Constituent Genera/Species Stone Quarry Porifera Demospongea Belemnospongia fascicularis Lower Unidentifiable tabulate coral? Lower Anthozoa Zaphrentoides spinulosus Absent Cnidaria Sphenothallus sp. Lower/ Intermediate Scyphozoa? Paraconularia missouriensis Passive Predator? Mollusca Pelecypoda Sulcatopinna missouriensis Intermediate Annelida? Polychaeta? Crinicaminus haneyensis Lower

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Table 6.6 (Continued)

Amphissites? Absent Bairdia? Arthropoda Ostracoda Absent Coronakirkbya? Absent Anthracospirifer leidyi Lower Cleiothyridina sublamellosa Lower Composita lewisensis Lower Composita subquadrata Lower Diaphragmus elegans Lower Articulata Dielasma sp. Lower Eumetria costata Lower Brachiopoda Eumetria verneuliana Lower Orthotetes kaskaskiensis Lower Punctospirifer sp. Lower Streptorhynchus sp. Lower Lingulipora sp. Lower Inarticulata Oehlertella pleurites Absent Orbiculoidea sp. Lower Anisotrypa sp. Lower Archimedes cf. meekanus Intermediate Eridopora sp. Lower Fenestella sp. Lower Fenestrellina sp. Lower Glyptopora punctipora Absent Lyroporella sp. Absent Bryozoa Stenolaemata Polypora sp. Lower Rhombopora sp. Lower Septopora sp. Lower Sulcoretepora sp. Absent Thamniscus? Lower/ Intermediate Unidentifiable encrusting bryozoan Lower

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Table 6.6 (Continued)

Unidentifiable edrioasteroid Lower Edrioasteroidea Lepidodiscus laudoni Lower Blastoidea Pentremities elegans Intermediate Lepidesthes formosa Absent Echinoidea Archaeocidaris megastylus Lower Ophiuroidea Unidentifiable encrinasterid Lower Acrocrinus shumardi Absent Agassizocrinus lobatus Lower Anartiocrinus maxvillensis Upper Aphelecrinus mundus Upper Camptocrinus cirrifer Intermediate Cryphiocrinus girtyi Intermediate Cymbiocrinus grandis Intermediate Echinodermata Cymbiocrinus tumidus Intermediate Dasciocrinus florealis Intermediate Eupachycrinus boydii Intermediate Crinoidea Linocrinus wachsmuthi Intermediate Pentaramicrinus gracilis Intermediate Phacelocrinus longidactylus Intermediate Phanocrinus bimagnaramus Intermediate Phanocrinus maniformis Intermediate Platycrinites sp. Intermediate Pterotocrinus acutus Absent Talarocrinus sp. Intermediate Taxocrinus whitfieldi Intermediate Tholocrinus spinosus Intermediate Zeacrinites magnoliaeformis Absent

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Table 6.6 (Continued)

Agassizodus Nektic Cladodus Nektic Ctenacanthus Nektic Deltodopsis Nektic Deltodus Nektic Chondrichthyes Listracanthus? Nektic (teeth) Lophodus Nektic Chordata Mesodmodus Nektic Psammodus? Nektic Polyrhizodus Nektic Taeniodus? Nektic Venustodus Nektic Chondrichthyes (dermal skin Petrodus Nektic plates)

Specimens from the coarse-grained calcarenite lithofacies (Figures 3.7, 6.2, 6.5 and 6.6), interpreted above to represent a high-energy shoal environment (Figures 6.2 and

6.3), appear to be randomly oriented and locally superimposed atop each other on stylolitic bedding-plane surfaces with no intervening shale partings (Figures 6.2, 6.4 and

6.5). Although disarticulated brachiopod valves are present, so are fully articulated valves, and crinoid calyces are preserved intact but lacking any stem segments (Figure

6.5). Slabs cut through some of the calcarenite layers show that some of the crinoid calyces were rapidly buried in migrating dunes (crossbedded) with their arms still flailing in the mobile sands (Figure 6.17). Blastoids are only preserved as isolated thecae, which probably readily rolled in the currents; stems and brachioles are always absent (Figure

6.18). The size-frequency distribution for Phanocrinus bimagnaramus (Figure 6.19) show that adults predominated, suggesting that the smaller juveniles may have been preferentially winnowed and transported elsewhere. Although 39 species are recorded 201

from this facies (Table 4.6), only three species, the brachiopod Anthracospirifer leidyi, the sponge Belemnospongia fascicularis, and the crinoid Phanocrinus bimagnaramus, were common on the shoals (Tables 4.6, 4.14, 5.3 and 6.2; Figure 6.20). The shoal communities probably formed on carbonate firm grounds that developed in low places on the shoals. However, during storms when these shoals were actively moving, community organisms were apparently ripped up, transported and buried by migrating sands. Hence, the stylolitic bedding planes may actually represent firm grounds on which some of the organisms lived as well as dumping grounds for others that had been transported.

Figure 6.17. Photo of slab cut through a crossbedded calcarenite from the shoal environment, showing two silicified crinoid calyces preserved along a stylolite surface with their arms still flailing in the mobile sands.

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Figure 6.18. Blastoid preserved as isolated thecae from the Valley Stone Quarry.

Valley Stone Quarry, Shoal: Histogram of Phanocrinus bimagnaramus Cup Width (cm) Normal

12 Mean 0.892 StDev 0.3121 N 25 10

u 6

F 4

0 0.3 0.6 0.9 1.2 1.5 Cup Width (cm)

Figure 6.19. Size-frequency histogram of the cladid crinoid, Phanocrinus bimagnaramus, cup width from the Valley Stone Quarry shoal community. The red curve shows how a normal distribution for the collection of specimens would appear.

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Figure 6.20. Schematic reconstruction of the interpreted, shallow, open-marine, Ramey Creek environmental continuum that developed seaward of the Tygarts Creek sandbelt, based on lithologies at the Valley Stone Quarry. Blue wavy line at top reflects sea level; the red dashed line reflects approximate normal wave base. Lithofacies here (Figure 6.3) are interpreted as depositional environments in color, and likely organism communities are shown as inset figures at the top. Organism rank and abundance in the inset figures are based on species frequency in Table 5.3. Organisms shown in the lower panel are drawn to a larger scale than the environments in which they lived.

The coarse-grained, argillaceous calcarenite lithofacies, interpreted to represent shoal-margin environments, contains more muddy sediments and exhibits shale partings

(Figures 6.2 and 6.3). Dissociated brachiopod valves and large fossil fragments characterize this lithofacies (Figure 6.21) and probably represent fossil debris transported off the shoal tops, whereas the mud in the lithofacies was probably winnowed from the shoals and sedimented in the quieter, deeper waters. Fossil preservation in this setting is not as good as that in the shoal setting, suggesting further transportation and comminution of the specimens. The size-frequency diagram for the Phanocrinus bimagnaramus in this setting shows a predominance of juveniles (Figure 6.22) that may represent some of those winnowed from the nearby shoals. Organism communities were probably present locally (Figure 6.20), but it is difficult to distinguish those from the materials that were carried in from the adjacent shoal setting. Twenty-nine species were recorded from this facies (Table 4.7), but the most abundant of those include the brachiopods Composita subquadrata and Anthracospirifer leidyi and the crinoid

Phanocrinus bimagnaramus (Tables 5.3 and 6.2).

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Figure 6.21. Appearance of a bedding plane from the fine-grained calcarenite and interbedded calcilutite lithofacies (shoal-margin environment) showing some of the fossil fragments that characterize the environment. Large shark tooth in the center (Cladodus) was probably transported into the area.

Valley Stone Quarry, Shoal-margin: Histogram of Phanocrinus bimagnaramus Cup Width (cm) Normal

8 Mean 0.6389 StDev 0.2810 7 N 18

u 4

r F 3

0 0.0 0.3 0.6 0.9 1.2 Cup Width (cm)

Figure 6.22. Size-frequency histogram of the cladid crinoid, Phanocrinus bimagnaramus, cup width for the Valley Stone Quarry shoal-margin community. The red curve shows how a normal distribution for the collection of specimens would appear.

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The fine-grained calcarenite and interbedded calcilutite lithofacies represents a deeper, transitional environment (Figures 6.2, 6.3 and 6.20). The typical preservation here reflects hardgrounds that were suddenly buried below shales (Figure 3.9). The hardgrounds contain attached edrioasteroids and most of the ophiuroids appear to be dorsal-side up, as if they were buried in place. Corrosion pits on the surface of the bed may reflect localized dissolution during exposure on the sea bottom (Figure 3.9).

Crinoids are typically preserved with stem segments intact, and the size-frequency distribution for Phanocrinus bimagnaramus is more normal (Figure 6.23), suggesting that the crinoids may have been living in place nearby. Twenty-one species are reported from this facies (Tables 4.8, 4.14 and 5.3), but the most common species are again the brachiopods Anthracospirifer leidyi and Composita subquadrata and the crinoid

Phanocrinus bimagnaramus (Tables 5.3 and 6.2). The setting for this environment was still close enough to the shoals that skeletal sands could be transported in during storms

(Figure 3.10), but far enough away that this did not happen often, so that these sands were exposed long enough that in-place lithification and corrosion to hardgrounds could occur (Figure 3.9). Most of the communities apparently developed on these hardgrounds, which were periodically buried during mud fallouts from nearby storms.

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Valley Stone Quarry, Transitional: Histogram of Phanocrinus bimagnaramus Cup Width (cm) Normal

Mean 0.645 14 StDev 0.3236 N 20 12

n 8

e r 6

0 0.2 0.6 1.0 1.4 Cup Width (cm)

Figure 6.23. Size-frequency histogram of the cladid crinoid, Phanocrinus bimagnaramus, cup width for the Valley Stone Quarry transitional community. The red curve shows how a normal distribution for the collection of specimens would appear.

The argillaceous calcilutite lithofacies (Figures 3.10 and 6.2) represents the deepest or basinal environments (Figure 6.3 and 6.20). Deeper environments like this are typically the most stable, and hence, have the greatest diversity and density as is indicated here in Tables 5.4, 5.5 and 5.7. The size-frequency distribution shows a slightly skewed normal distribution, but with contributions from all size ranges, suggesting presence of communities (Figure 6.24). Specimens from this environment were typically buried rapidly in mud fallouts and exhibit the best preservation of any of the Valley Stone lithofacies; delicate calyces, thecae, plates, arms, pinnules and spines from echinoids and crinoids are preserved nearly intact (Figures 1.2, 3.3, 3.10, 6.25 and 6.26).

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Valley Stone Quarry, Basinal: Histogram of Phanocrinus bimagnaramus Cup Width (cm) Normal

Mean 0.8333 5 StDev 0.3939 N 12

c 3

F 2

0 0.2 0.6 1.0 1.4 Cup Width (cm)

Figure 6.24. Size-frequency histogram of the cladid crinoid, Phanocrinus bimagnaramus, cup width for the Valley Stone Quarry basinal community. The red curve shows how a normal distribution for the collection of specimens would appear.

Nonetheless, the presence of splayed arms (Figure 6.25) and thin graded beds

(Figure 6.27) indicate that energetic currents and storms probably did occasionally reach this environment. Forty-nine species are recorded from this lithofacies, making it the richest of the fossil communities in this study (Tables 5.4, 5.5 and 5.7). Composita subquadrata, Belemnospongia fascicularis, and Pentremites elegans were the most abundant invertebrates. However, chondrichthyan fragments are also relatively common in the community, probably reflecting transportation from outside sources.

Biostratinomic and sedimentologic indicators from the Valley Stone Quarry indicate that the fossils found there represent “transported fossil assemblages,” using the terminology of Fagerstrom (1964). Nonetheless, based on the observations of Kidwell

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and Flessa (1996) and Kidwell (2001), it is likely that these assemblages approximate actual communities from the past. Different types of preservation in the lithofacies apparently represent different energy regimes and sedimentologic processes, such that the deeper water, basinal and transitional environments, exhibit the best presentation (Figures

1.2, 3.3, 3.9, 6.25 and 6.26). Surprisingly, the shoal environment shows the next best preservation (Figures 6.5 and 6.6), largely because the communities were rapidly buried beneath migrating sand. In contrast, the shoal-margin environment shows the worst fossil preservation (Figures 3.8 and 6.21) with largely fragmented and dissociated fossil remains. Fossils preserved in this setting were apparently reworked on the shoal and subsequently into the shoal-margin setting, probably in large part by storm-related backflow currents. Because of the different communities, depositional environments and the processes active in each, the Valley Stone Quarry environments are not as easily interpreted as those in the 213 Quarry, but interpretations of how the environments may have appeared are presented in Figure 6.20.

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Figure 6.25. Detailed preservation of the cladid crinoid, Aphelocrinus mundus, preserved in argillaceous calcilutite from the basinal environment. The arms have been splayed outward to expose the anal chimney.

Figure 6.26. Bottom of the bed showing two Agassizocrinus lobatus specimens in apparent life position from the argillaceous calcilutite lithofacies, having been buried in argillaceous, micritic mud. Note the detailed preservation of pinnules on the arms.

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Figure 6.27. Thin graded bed from the argillaceous calcilutite lithofacies. Centimeter scale to the right.

6.4. Interpretation of relative ordinal and statistical data

In my attempt to characterize and compare fossils at the two collecting localities,

5,137 specimens from 15 classes and 77 species were analyzed for taxonomy and relative abundance (Tables 4.12–4.14; Figures 5.1–5.50). The large numbers of specimens analyzed, probably the largest, megascopic, fossil communities ever analyzed from this time, necessarily require the use of some type of statistical analyses for comparison.

Hence, in this section, I will first examine implications of the taxonomic differences between communities based on the simple ordinal counting and comparison of taxonomic groups (classes, genera or species) collected in Chapter 4 and follow this with interpretations based on the statistical measures of abundance from Chapter 5.

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6.4.1. The 213 Quarry

As already indicated, the early collecting at both localities was done to characterize what were thought to be “crinoid gardens.” However, the work in this study immediately showed that other organisms were far more abundant, and perhaps more important, that the crinoids. At the 213 Quarry, the most abundant organisms in both communities were bryozoans (Archimedes and rhomboporid bryozoans; Tables 4.12 and

5.2), comprising almost 67% of the population. Bryozoans were followed in abundance by six species of brachiopods at 15%, of which only one species (Anthracospirifer leidyi at 67% of the brachiopods) predominated. Sixteen species of crinoids were present for nearly 7%, of which one species predominated (Cymbiocrinus grandis at 23% of the crinoids). Clearly, this community was not a “crinoid garden,” but rather a “bryozoan thicket,” in which two non-echinoderm species, Archimedes cf. meekanus and

Anthracospirifer leidyi, predominated (Table 6.2). All of these organisms were filter feeders (Tables 6.3 and 6.4), but the dominant forms apparently partitioned the filter- feeding niche into lower (rhomboporid bryozoans, brachiopods), intermediate

(Archimedes and some crinoids) and upper tiers (some crinoids), perhaps to lessen competition. Browsers and deposit feeders (echinoids, ostracods), passive predators

(corals, polychaetes, and some crinoids), and scavengers (echinoids, ostracods) were also present, but in very low numbers (<1%).

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locality. Organism density is provided in Table 5.7. Species diversity and evenness are then interpreted using an adapted version of Pearson’s “r” (Table 1.2).

Table 6.7 Summary for the 213 and Valley Stone quarries, species richness, diversity, evenness, and interpretation based on an adaptation of Pearson’s “r”.

Interpretation of Simpson’s Interpretation Quarry, Simpson’s Shannon’s Index of of Shannon’s Species Location/Lithologies Index of Evenness Diversity, Evenness Richness (facies) Diversity Index using Index, using Pearson’s “r” Pearson’s “r” (Table 1.2) (Table 1.2) Somewhat Entire 213 Quarry Moderate Strong Landscape 45 0.49 0.68 Diversity Evenness Somewhat strong Moderate 213 – Top-of-slab 44 0.63 0.46 Diversity Evenness Somewhat Somewhat Strong Strong 213 – Bottom-of-slab 25 0.79 0.61 Diversity Evenness Entire Valley Stone Somewhat Quarry (VS) Strong Strong Landscape 65 0.90 0.69 Diversity Evenness Somewhat VS – Coarse-grained Strong Strong calcarenite (Shoal) 39 0.89 0.71 Diversity Evenness VS – Coarse-grained, argillaceous Somewhat calcarenites (Shoal- Strong Moderate margin) 29 0.66 0.57 Diversity Evenness VS – Fine-grained calcarenites and interbedded calcilutites Strong Strong (Transitional) 21 0.90 0.84 Diversity Evenness Somewhat VS – Argillaceous Strong Strong calcilutites (Basinal) 49 0.92 0.76 Diversity Evenness

Examining the 213 Quarry, the highest species richness (45) and density (798 indiv./m2) were found in the top-of-slab community, whereas the bottom-of-slab community contains nearly one-third fewer species (25) across the same area for a density of 200 indiv./m2, a nearly 75% decrease in individuals per m2. Moreover, because 214

two species (Archimedes cf. meekanus and rhomboporid bryozoans) comprise nearly

70% of the top-of-slab community, the evenness is lower than for the bottom-of-slab community, where the two top species (Archimedes cf. meekanus and rhomboporid bryozoans) comprise only 60% of the community. Hence, the bottom-of-slab community has a more even distribution of species (0.69) compared to the top-of-slab community

(0.46), and when the difference in evenness is considered, the diversity index for the bottom-of-slab community (0.79) is about 20% larger than that of the top-of slab community (0.63). Hence, even though the top-of-slab community has greater species richness and the bottom-of-slab community has greater species evenness, there is a trade off in values in the diversity index such that the diversity indices for both are interpreted as “somewhat strong” using Pearson’s “r” (Table 6.7). Looking at the two communities together or as a “landscape” with 45 species, that landscape exhibits a moderate diversity and somewhat strong evenness (Table 6.7). That richness and diversity are not greater, may reflect the fact that the area of community development (Figure 6.1) was subject to frequent disruption by storm activity, or possibly, that the studied area was relatively small.

6.4.2. Valley Stone Quarry

At the Valley Stone Quarry as a whole, two brachiopods, Composita subquadrata and Anthracospirifer leidyi, are generally the predominant species, comprising nearly

36% of the fauna, followed by the sponge Belemnospongia fascicularis at about 12%.

Eighteen species of crinoids are present at the Valley Stone Quarry, comprising about

24% of the entire landscape community. One crinoid species, Phanocrinus

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bimagnaramus, predominates in every community and includes 31% of all crinoids present. One or the other of these two brachiopod species is the dominant species in each of the interpreted environments, but in three of the environments a sponge or crinoid may rank second or third in abundance. As in the 213 Quarry, most of the organisms were filter feeders (Tables 6.3 and 6.4), but the dominant forms apparently partitioned the filter-feeding niche into lower (brachiopods, sponges and some bryozoan), intermediate

(Archimedes, Pentremites and some crinoids) and upper tiers (some crinoids). Browsers and deposit feeders (echinoids, ophiuroids), passive predators (corals, polychaetes, and some crinoids), and scavengers (echinoids, ophiuroids) were also present, but in very low numbers (<1%).

In the coarse-grained calcarenite lithofacies (shoal environment), two species, the brachiopod Anthracospirifer leidyi and the sponge Belemnospongia fascicularis comprise about 42% of the shoal community followed by the crinoid Phanocrinus bimagnaramus at about 9% (Figure 6.20; Table 6.2). On the shoal-margin community, one brachiopod,

Composita subquadrata predominates at more than 57%, whereas the next most abundant species, Anthracospirifer leidyi and Phanocrinus bimagnaramus comprise only 13% of the community (Figure 6.20; Table 6.2). In the transitional community, Anthracospirifer leidyi predominates at about 20%, whereas the next two species in abundance

Phanocrinus bimagnaramus and Phanocrinus bimagnaramus each comprise slightly more than 13% of their communities (Figure 6.20; Table 6.2). In contrast, in the basinal community, the three most abundant fauna, Composita subquadrata (17.5%),

Belemnospongia fascicularis (12%) and Pentremites elegans (8.2%), comprise only 38%, meaning that several other species are present but at lower numbers. The densities of

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individuals in the shoal, shoal-margin, and transitional environments are not that different from each other, averaging about 370 individuals/m2, but are about three times less dense than the basinal community, which, at about 1024 individuals/m2, reflects the highest density of organisms in any community in this study.

Using the data generated in Chapters 4 and 5, as well as Appendix A, Table 6.7 provides a summary of the statistical measures of abundance in the form of species richness (Table 5.4), diversity (Table 5.5) and evenness (Table 5.6) for each of the lithofacies (environments) examined in this study, as well for the entire landscape at each locality. Organism density is provided in Table 5.7. Species diversity and evenness are then interpreted using an adapted version of Pearson’s “r” (Table 1.2). In the Valley

Stone Quarry, the highest species richness (49) and density (1024 indiv./m2) were found in the basinal community, whereas the transitional community has the lowest richness

(21) and density (320 indiv./m2), contains nearly one-half fewer species (21) at nearly one-third the density of the basinal community. In terms of evenness, the shoal, transitional and basinal communities exhibit somewhat strong to strong evenness (0.71–

0.84), with the shoal-margin community showing the lowest evenness (0.57), because of the dominance of Composita subquadrata at 57% of the community. The dominance of

Composita subquadrata in the shoal-margin community is also reflected in the diversity indices such that the shoal- margin community exhibits a diversity (0.66) that is about

27% less than the diversity of the other environments, which average about 0.90.

Examining all of the Valley Stone Quarry communities together or as a “landscape,” shows a total of 65 species that exhibits strong diversity and somewhat strong evenness

(Table 6.7). It is likely that the high species richness (65) and strong diversity (0.90)

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across the entire quarry (landscape) reflect a greater number of habitats in which organisms can live and the fact that some of the environments are deeper and more stable.

The lower organism density (465 indiv./m2), however, probably reflects the fact that the organisms are spread across a much larger area. Is is important to note that the richness, diversity and density figures noted above may also reflect the larger number of samples and multiple beds that were available for study at this locality.

6.4.3. Locality comparisons

At the start, it is important to indicate that comparisons between the two localities are not necessarily as balanced as one could hope for, because the localities reflect vastly different areas and depositional environments. Although both localities reflect deposition at nearly the same time, the 213 Quarry exhibits two communities (bottom- and top-of- bed) that developed successively in one small area of about 3.3 m2 (35 ft2), whereas the

Valley Stone Quarry reflects at least four communities that developed across a larger area of approximately 0.17 km2 (0.07 mi2) in four environments. Hence, although I will make comparisons for entire landscapes (all environments) at the two localities, comparison of the individual environments at each locality is probably more accurate and balanced.

6.4.3.1. Depositional environments

Using the term “landscape” in the sense of Patzkowsky and Holland (2012), I describe the landscape and depositional environment in the Ramey Creek Member at the

213 Quarry (Figure 3.13) as a mid-ramp storm shelf (e.g., Aigner, 1982; Fichter and

Poché, 1993) (Figure 6.1) on the western margin of the Appalachian Basin. Having been on the southern downdropped block in east-central Kentucky, environment here was

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somewhat deeper (e.g., Ettensohn, 1975a, b, 1977, 1981a, b) than those at the Valley

Stone Quarry, but apparently not necessarily more stable because of frequent storm intrusion. In contrast, the Ramey Creek Member at the Valley Stone Quarry represents deposition on a structurally higher platform (Figure 6.1), wherein waves, tides and storm- related currents could rework sediments in a shallower setting across a broad area to generate several depositional environments (e.g., Ettensohn, 1975a, b, 1977, 1981a, b)

(Figure 6.3). Four recognized lithofacies have been interpreted to represent four more or less distinct depositional environments (Figures 6.2 and 6.3), in comparison to the one depositional environment at the 213 Quarry.

6.4.3.2. Comparing statistical parameters

Statistical parameters used for comparison include species richness, diversity, evenness, density, and levels of diversity (α, β, γ), which can be found in Tables 5.4, 5.5,

5.6, 5.7, 5.10, 5.11, 5.12 and 6.7. Species richness, which is the same as α-diversity

(Whittaker, 1972; Patzkowsky and Holland, 2012) reflects the total number of species in an environment or locality. At the 213 Quarry, the top-of-slab community (S = 44) has nearly two times the species richness as the bottom-of-slab community (S = 25). The reason for this is uncertain, except for the possibility that the top-of slab community may have persisted longer before burial, and therefore, was able to recruit more species over time. The total species richness for the entire 213 landscape is 45 species. In contrast, the total species richness for the Valley Stone Quarry is 65, nearly 1.5 times greater than that of the 213 Quarry, probably reflecting the greater area of the Valley Stone landscape

(Figures 1.4 and 6.3). Of the four Valley Stone environments (Figure 6.3), the basinal and shoal have higher richness values (average = 45), comparable to the top-of-the-bed

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community at the 213 Quarry (44), whereas the shoal-margin and transitional environments have richness values (average = 25) that are comparable to the bottom-of slab-community (25) at the 213 Quarry (Table 6.7). The greater richness of the basinal and shoal environments may reflect the stability of deeper-water environments and the presence of higher energies and fast-lithifying substrates that would have attracted rheophilic organisms like crinoids, respectively. Shoal-margin and transitional environments may have experienced frequent burial, like the 213 storm shelf, from sediments washed off the shoals, thus limiting the number of species that could colonize the areas.

As for the evenness of the species in most of these communities, species evenness is relatively strong across all communities, except for the 213 top-of-slab community (EH

= 0.46) and the Valley Stone shoal-margin community (EH = 0.57), which show dominance by one species each, Archimedes cf. meekanus and Composita subquadrata, respectively (Tables 6.2 and 6.7). Why these two species became so dominant in their respective environments is uncertain, but their dominance may reflect the fact that their environments fulfilled specialized feeding needs for each. Evenness can also be expressed in rank-abundance graphs (Figures 5.51, 5.52 and 5.53).

Diversity (D), on the other hand, is clearly greater across the Valley Stone landscape (D = 0.90) than for the 213 Quarry landscape (D = 0.49). Nearly every environment in the Valley Stone landscape supported strong diversity, in contrast to the somewhat strong to moderate diversity in the 213 Quarry, which reflects community dominance by two bryozoan species (Table 6.2). Also, the larger area across which each

Valley Stone environment extends creates more habitat and niche possibilities for

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different organisms. Similarly, the larger area of the Valley Stone landscape and each of its environments is also indicated in the smaller densities of the Valley Stone landscape and most of its communities, compared to the 213 communities (Table 5.7). Although the

213 Quarry bottom-of-slab community has a relatively low density (~200 indiv./m2), it is dwarfed by the larger top-of-slab community with nearly five times the density (997 indiv./m2), which suggest that the organisms there were living in very crowded conditions in localized areas (Figure 6.14). The Valley Stone Quarry, in general, departs from this cramped state with more moderate densities that range from 320 to 419 indiv./m2, with the exception of the basinal environment, where densities reached 1,024 indiv./m2. These kinds of numbers probably reflect the increased stability of deeper-water environments and the fact fewer basinal environments were present (Figure 6.20).

Alpha diversity (α) has already been discussed in the form of species richness (α =

S) with the Valley Stone Quarry having the higher richness by 20 species (Tables 5.10 and 6.7). Beta diversity (β), in contrast, compares changes in the species richness of two communities or areas. Hence, the change in species richness between the 213 and Valley

Stone quarries is 44 (Table 5.10), meaning that 44 species do not occur in one or the other landscape. Most of these species are sporadic to rare (Tables 5.1, 5.2 and 5.3) in both landscapes (Tables 4.10, 4.12 and 4.13), but Belemnospongia fascicularis, which makes up 12% of the Valley Stone communities, is an exception. It apparently had broader environmental tolerances than any of the other β-diversity species did. In comparing species occurrence between other environments, the β-diversity ranges from

18 to 36, often with increasing β-diversity with community depth (Table 5.12), meaning

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that more different organisms enter successively deeper environments, probably reflecting the environmental stability that comes with greater depth.

The final measure of diversity is gamma diversity (γ), which measures the overall diversity for the different ecosystems within a region (Hunter, 2002). Assuming that all environments in the 213 and Valley Stone quarries reflect their own regional settings, then their γ-diversities are 45 and 65, respectively. However, if I assume that the communities in both quarries reflect a larger, eastern Kentucky or western Appalachian

Basin region, then the γ-diversity is 77 (Tables 5.10, 5.11, and 5.12).

6.4.3.3. Comparing dominant species

The dominant species in the 213 Quarry landscape is Archimedes cf. meekanus followed by rhomboporid bryozoans (Table 6.2) both of which are low- to mid-level filter feeders and contain delicate parts that cannot withstand high energies. Crinoids are the dominant echinoderms in the community, but compared to the bryozoans and brachiopods, they are either sporadic or rare, comprising only about 7% of the communities. In the Valley Stone Quarry landscape, the dominant species is the brachiopod Composita subquadrata followed by another brachiopod Anthracospirifer leidyi and the sponge Belemnospongia fascicularis (Table 6.2), all of which are low-level filter feeders. Crinoids are the dominant echinoderms and make up about 24% of the communities, more than three times that of the crinoids at the 213 Quarry localities.

The abundance of delicately constructed, mid-level-feeding bryozoans in the 213

Quarry environments suggests less energetic, deeper, mid-ramp environments. The presence of mid- and high-level feeding crinoids suggests that currents were present, as

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most crinoids are rheophilic feeders (Ausich and Bottjer, 1982; Bottjer and Ausich,

1986), but these currents were apparently not strong enough to destroy the delicate bryozoans. Moreover, the crinoids may have further aided bryozoan and other low-tier feeding by baffling currents that passed by. These kinds of deeper-water settings would have been common during Ramey Creek Member deposition on the southern- downdropped block (Figures 2.4 and 6.1). However, when storms crossed the area, very high-energy currents and waves would have buffeted the area, knocking down delicate structures and ripping up stemmed organisms, only to be buried later in the accompanying sediment fallout.

The dominance of small, sturdy, low-level filter-feeding brachiopods and sponges in the Valley Stone Quarry contrasts markedly with the delicate bryozoans that predominate at the 213 Quarry and suggests an overall shallower area, prone to high- energy. Not only were these environments shallower, but they were also apparently closer to wave base where waves frequently reworked and winnowed fines from the skeletal sands. The low profiles of the most common organisms would have made them less susceptible to the effects of agitation. So higher energies seem to have been a common denominator at the Valley Stone Quarry locality, and this may well be related to an uplifted, platform-like setting on the northern uplifted block (Figures 2.4 and 6.1). This same setting may also account for the more varied environmental setting of the Ramey

Creek Member in the Valley Stone Quarry (Figure 6.20) and the generally lesser organism density. Interestingly, the shoal environment at the Valley Stone Quarry exhibits more crinoid individuals (nearly 50% of all Valley Stone environments) than any other environment, perhaps reflecting the generally rheophilic nature of crinoids.

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However, most of the crinoids belong to one species, Phanocrinus bimagnaramus, whose unusual arm structure, with two extremely thick arms protecting the more delicate inner arms (Figure 6.28), may be an adaptation to higher energies. Storms, no doubt, crossed this area as well, burying communities across the entire landscape.

Figure 6.28. Photo of Phanocrinus bimagnaramus, whose unusual arm structure, with two extremely thick arms protecting the more delicate inner arms. This kind of arm structure may be an adaptation to living and feeding in high-energy environments.

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CHAPTER 7: CONCLUSIONS

The purpose of this study was to characterize two Late Mississippian fossil echinoderm communities from the Ramey Creek Member of the Slade Formation in eastern Kentucky and try to understand the differences between them. These communities not only contained nearly 900 echinoderms, but more than 4,100 non-echinoderm organisms, and hence, provide an opportunity understand broad community relationships.

This is not the first study of this kind (e.g., Brett and Taylor, 1997), but with more than

5,000 megascopic specimens, it is at present the largest of its kind and one of only two studies of Late Mississippian (Chesterian) echinoderm communities. Why are echinoderm communities so significant? Although relatively common organisms in today’s seas and oceans, their preservation potential is very limited because they are composed of many ossicles or plates that rapidly disaggregate after death, meaning that most do not enter the fossil record. Hence, they are rare as fossils and information about how they lived in the past has been difficult to acquire, and a major goal of this project is to provide more complete regional and local contexts in which to understand echinoderms and their roles in community organization. The major conclusions of this study are listed below.

1. Previous work in eastern Kentucky suggests that the Kentucky River Fault was

apparently active during Mississippian deposition, such that the northern block

was probably uplifted relative to the southern block and experienced shallower,

more energetic conditions. However, the possibility that differences between the

two localities merely reflect normal facies variability cannot be precluded.

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2. The Ramey Creek Member at both localities represents a shallow, open-marine

environment that is manifest in different ways at the localities. Some of the

differences, however, may reflect the number and size of samples at each locality.

The 213 Quarry locality reflects a storm-shelf setting characterized by a

tempestite lithofacies with two somewhat similar communities: top-of-slab and

bottom-of slab communities. The Valley Stone Quarry, in contrast, represents an

uplifted platformal area that differentiated into four different lithofacies: a coarse-

grained calcarenite lithofacies that indicates a shoal environment; coarse-grained

argillaceous calcarenite lithofacies indicating a shoal-margin environment; fine-

grained calcarenite and interbedded calcilutite lithofacies representing a

transitional environment; and an argillaceous calcilutite lithofacies reflecting a

basinal environment.

3. The collection at the 213 Quarry represents a relatively small, isolated

community, which may reflect the limited sample size. In contrast, the collection

at the Valley Stone Quarry represents a broader continuum of four environments,

each with its own community. The larger platformal area and shallower waters

help to explain the larger number of lithofacies and communities at the Valley

Stone Quarry locality.

4. Biostratinomic indicators, such as faunal composition, fossil density, size-

frequency distribution, dissociation of hard parts, surface conditions, and fossil

orientation were examined to develop a brief history of each community.

Integrating these indicators suggests that most communities were ripped up in

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high-energy events, transported short distances, and exposed briefly, only to be

buried rapidly by storm-related sediments or migrating sand dunes.

5. An adaptation of the statistical parameter, Pearson’s “r,” was newly adapted and

applied for the descriptive interpretation of diversity and evenness indices.

6. The 213 Quarry community was dominated by lower- and mid-level filter feeders,

mainly bryozoans; the echinoderms, mainly crinoids, were minor constituents of

the communities at 7%. In general, this community exhibited lower species

richness, lower diversity, high density and moderate evenness than the Valley

Stone Quarry communities, mostly reflecting the overwhelming abundance of two

bryozoan species at 67% of the community. The abundance and delicate nature of

the predominant bryozoans suggests that the original communities developed in

quiet, deeper waters. The 213 Quarry community was very localized in extent

along the storm shelf, but this situation may reflect the limited sampling area.

7. The Valley Stone Quarry communities were dominated by lower-level filter

feeders, namely brachiopods and sponges; echinoderms, mainly crinoids, were

only modest constituents of the communities at 24%. In general, the Valley Stone

Quarry communities exhibited higher species richness, higher diversity, lower

density, and higher evenness, reflecting the greater number of available

environments and larger community areas, or the larger sampling area. The most

abundant species, brachiopods, comprised only 40% of the communities, and the

small, lower-profile and sturdy nature of the dominant organisms suggests a

higher-energy setting, and this is supported by the dominance of rheophilic

crinoids.

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8. Most of the organisms found in these communities have been previously

described and are know well enough to identify despite silicification. However,

one new species of the ophiuroid Schoenaster was discovered and will be

formally described in a separate publication.

9. The α-diversity of the 213 and Valley Stone quarries was 45 and 65, respectively,

but the β-diversity, or change in species between the localities, was 44, mostly

reflecting different species at the Valley Stone locality, which again may reflect

the larger sampling size. The total diversity across the entire eastern Kentucky

area, or γ-diversity, was 77 species, reflecting five echinoderm classes, and 10

non-echinoderm classes.

10. Although the fossil faunas were originally collected for their echinoderm-

dominated “crinoid gardens,” community analyses indicate that echinoderms were

only minor to modest constituents of these communities, and that bryozoans,

brachiopod and sponges were far more abundant. In effect, the 213 Quarry

communities were “bryozoan thickets,” and the Valley Stone Quarry communities

were “brachiopod pavements.” Most of the echinoderms were probably present in

response to the ambient energy regimes, but as higher-level bafflers may have

enhanced the environment for lower-level feeders. Although storm- and wave-

related events appear to have preserved the communities, community makeup

appears to have been largely controlled by depth and energy levels possibly

related to larger-scale, structural controls or to variations in lateral facies

distribution.

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APPENDIX A

STATISTICAL DATA FOR FOSSILS FROM BOTH QUARRIES

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Table TA1. Simpson’s Index of Diversity and Shannon’s Evenness calculations for the 213 Quarry entire landscape.

Proportion of entire Natural Log 213 Quarry Entire Landscape Number of population of - Simpson's Index of Diversity Individuals to Proportion and Shannon's Evenness individual Calculations species Number P * ln P * - n(n-1) P ln P (n) 1

Anthozoa (1) Zaphrentoides spinulosus 7 42 0.002158495 -6.138343957 0.013249586

230 Scyphozoa? (1)

Sphenothallus sp. 8 56 0.002466852 -6.004812565 0.014812982

Pelecypoda (1) Sulcatopinna missouriensis 1 0 0.000308356 -8.084254106 0.002492832

Polychaeta? (1) Crinicaminus haneyensis 1 0 0.000308356 -8.084254106 0.002492832

Ostracoda (3) Amphissites? 1 0 0.000308356 -8.084254106 0.002492832 Bairdia? 1 0 0.000308356 -8.084254106 0.002492832 Coronakirkbya? 3 6 0.000925069 -6.985641818 0.006462203

Table TA1. (Continued)

Articulata (6) Anthracospirifer leidyi 316 99540 0.097440641 -2.328511893 0.226891692 Cleiothyridina sublamellosa 55 2970 0.016959605 -4.076920921 0.069142970 Composita lewisensis 19 342 0.005858773 -5.139815127 0.030113009 Composita subquadrata 65 4160 0.020043170 -3.909866836 0.078366125 Diaphragmus elegans 23 506 0.007092199 -4.948759890 0.035097588 Eumetria verneuliana 3 6 0.000925069 -6.985641818 0.006462203

Inarticulata (2) Oehlertella pleurites 2 2 0.000616713 -7.391106926 0.004558191 Orbiculoidea sp. 8 56 0.002466852 -6.004812565 0.014812982

231 Stenolaemata (9)

Archimedes cf. meekanus 1738 3018906 0.535923528 -0.623763800 0.334289696 Eridopora sp. 16 240 0.004933703 -5.311665384 0.026206181 Fenestella sp. 12 132 0.003700278 -5.599347457 0.020719140 Glyptopora punctipora 135 18090 0.041628122 -3.178979328 0.132334940 Lyroporella sp. 1 0 0.000308356 -8.084254106 0.002492832 Polypora sp. 99 9702 0.030527290 -3.489134256 0.106513812 Rhombopora sp. 432 186192 0.133209991 -2.015828518 0.268528498 Septopora sp. 8 56 0.002466852 -6.004812565 0.014812982 Sulcoretepora sp. 4 12 0.001233426 -6.697959745 0.008261437

Blastoidea (1) Pentremities elegans 41 1640 0.012642615 -4.37068204 0.055256850

Echinoidea (2) Lepidesthes formosa 1 0 0.000308356 -8.084254106 0.002492832

Table TA1. (Continued)

Archaeocidaris megastylus 3 6 0.000925069 -6.985641818 0.006462203 (isolated spines only)

Crinoidea (16) Acrocrinus shumardi 1 0 0.000308356 -8.084254106 0.002492832 Agassizocrinus lobatus 1 0 0.000308356 -8.084254106 0.002492832 (juvenile) Anartiocrinus maxvillensis 1 0 0.000308356 -8.084254106 0.002492832 Aphelecrinus mundus 5 20 0.001541782 -6.474816194 0.009982757 Camptocrinus cirrifer 12 132 0.003700278 -5.599347457 0.020719140 Cryphiocrinus girtyi 2 2 0.000616713 -7.391106926 0.004558191 Cymbiocrinus grandis 55 2970 0.016959605 -4.076920921 0.069142970 Cymbiocrinus tumidus 11 110 0.003391921 -5.686358834 0.019287680

232 Dasciocrinus florealis 35 1190 0.010792476 -4.528906045 0.048878110

Pentaramicrinus gracilis 28 756 0.008633981 -4.752049596 0.041029105 Phanocrinus bimagnaramus 29 812 0.008942337 -4.716958276 0.042180632 Phanocrinus maniformis 4 12 0.001233426 -6.697959745 0.008261437 Pterotocrinus acutus 16 240 0.004933703 -5.311665384 0.026206181 Taxocrinus whitfieldi 31 930 0.009559050 -4.650266902 0.044452135 Tholocrinus spinosus 4 12 0.001233426 -6.697959745 0.008261437 Zeacrinites magnoliaeformis 3 6 0.000925069 -6.985641818 0.006462203

Chondrichthyes (2) Cladodus 1 0 0.000308356 -8.084254106 0.002492832 Venustodus 1 0 0.000308356 -8.084254106 0.002492832

Grand Total: 3,243 3,349,854 1.8501984

Table TA2. Simpson’s Index of Diversity and Shannon’s Evenness calculations for the Valley Stone Quarry entire landscape.

Proportion of entire Natural Valley Stone Quarry entire Number of population Log of landscape - Simpson's Index of Individuals to Proportion Diversity and Shannon's individual Evenness Calculations species Number P * ln P * - n(n-1) P ln P (n) 1

Demospongea (1) Belemnospongia fascicularis 233 54056 0.123020063 -2.0954078 0.257777203

233 Anthozoa (1) Unidentifiable tabulate coral? 1 0 0.000527983 -7.5464463 0.003984396

Scyphozoa? (2) Sphenothallus sp. 28 756 0.014783527 -4.2142418 0.062301357 Paraconularia missouriensis 8 56 0.004223865 -5.4670047 0.023091889

Pelecypoda (1) Sulcatopinna missouriensis 4 12 0.002111932 -6.1601519 0.013009825

Table TA2. (Continued)

Polychaeta? (1) Crinicaminus haneyensis 2 2 0.001055966 -6.8532991 0.007236852

Articulata (11) Anthracospirifer leidyi 272 73712 0.143611404 -1.9406442 0.278698640 Cleiothyridina sublamellosa 8 56 0.004223865 -5.4670047 0.023091889 Composita lewisensis 8 56 0.004223865 -5.4670047 0.023091889 Composita subquadrata 410 167690 0.216473073 -1.5302891 0.331266387 Diaphragmus elegans 26 650 0.013727561 -4.2883497 0.058868581 Dielasma sp. 1 0 0.000527983 -7.5464463 0.003984396 Eumetria costata 4 12 0.002111932 -6.1601519 0.013009825 Eumetria verneuliana 5 20 0.002639916 -5.9370084 0.015673201

234 Orthotetes kaskaskiensis 7 42 0.003695882 -5.6005361 0.020698919

Punctospirifer sp. 7 42 0.003695882 -5.6005361 0.020698919 Streptorhynchus sp. 1 0 0.000527983 -7.5464463 0.003984396

Inarticulata (2) Lingulipora sp. 2 2 0.001055966 -6.8532991 0.007236852 Orbiculoidea sp. 6 30 0.003167899 -5.7546868 0.018230264

Stenolaemata (10) Anisotrypa sp. 3 6 0.001583949 -6.4478340 0.010213042 Archimedes cf. meekanus 17 272 0.008975713 -4.7132329 0.042304625 Eridopora sp. 1 0 0.000527983 -7.5464463 0.003984396 Fenestella sp. 4 12 0.002111932 -6.1601519 0.013009825 Fenestrellina sp. 1 0 0.000527983 -7.5464463 0.003984396 Polypora sp. 24 552 0.012671595 -4.3683924 0.055354498 Rhombopora sp. 45 1980 0.023759240 -3.7397838 0.088854419

Table TA2. (Continued)

Septopora sp. 1 0 0.000527983 -7.5464463 0.003984396 Thamniscus? 40 1560 0.021119324 -3.8575668 0.081469204 Unidentifiable encrusting bryozoan 6 30 0.003167899 -5.7546868 0.018230264

Edrioasteroidea (2) Unidentifiable edrioasteroid 6 30 0.003167899 -5.7546868 0.018230264 Lepidodiscus laudoni 1 0 0.000527983 -7.5464463 0.003984396

Blastoidea (1) Pentremities elegans 89 7832 0.046990496 -3.0578099 0.143688005

Echinoidea (1)

235 Archaeocidaris megastylus 54 2862 0.028511088 -3.5574622 0.101427117

Ophiuroidea (1) Unidentifiable encrinasterid brittle star 34 1122 0.017951426 -4.0200857 0.072166270

Crinoidea (18) Agassizocrinus lobatus 30 870 0.015839493 -4.1452489 0.065658641 Anartiocrinus maxvillensis 31 930 0.016367476 -4.1124591 0.067310576 Aphelecrinus mundus 25 600 0.013199578 -4.3275704 0.057122102 Camptocrinus cirrifer 2 2 0.001055966 -6.8532991 0.007236852 Cryphiocrinus girtyi 1 0 0.000527983 -7.5464463 0.003984396 Cymbiocrinus grandis 35 1190 0.018479409 -3.9910982 0.073753135 Cymbiocrinus tumidus 23 506 0.012143611 -4.4109521 0.053564888 Dasciocrinus florealis 88 7656 0.046462513 -3.0691095 0.142598539 Eupachycrinus boydii 1 0 0.000527983 -7.5464463 0.003984396

Table TA2. (Continued)

Linocrinus wachsmuthi 48 2256 0.025343189 -3.6752453 0.093142435 Pentaramicrinus gracilis 16 240 0.008447730 -4.7738576 0.040328258 Phanocrinus bimagnaramus 141 19740 0.074445618 -2.5976864 0.193386368 Phanocrinus maniformis 1 0 0.000527983 -7.5464463 0.003984396 Phacelocrinus longidactylus 1 0 0.000527983 -7.5464463 0.003984396 Platycrinites sp. 1 0 0.000527983 -7.5464463 0.003984396 Talarocrinus sp. 6 30 0.003167899 -5.7546868 0.018230264 Taxocrinus whitfieldi 11 110 0.005807814 -5.1485510 0.029901827 Tholocrinus spinosus 1 0 0.000527983 -7.5464463 0.003984396

Chondrichthyes (13) Agassizodus 30 870 0.015839493 -4.1452489 0.065658641

236 Cladodus 31 930 0.016367476 -4.1124591 0.067310576

Ctenacanthus 25 600 0.013199578 -4.3275704 0.057122102 Deltodopsis 2 2 0.001055966 -6.8532991 0.007236852 Deltodus 1 0 0.000527983 -7.5464463 0.003984396 Listracanthus? 35 1190 0.018479409 -3.9910982 0.073753135 Lophodus 23 506 0.012143611 -4.4109521 0.053564888 Mesodmodus 88 7656 0.046462513 -3.0691095 0.142598539 Petrodus 1 0 0.000527983 -7.5464463 0.003984396 Psammodus? 48 2256 0.025343189 -3.6752453 0.093142435 Polyrhizodus 16 240 0.008447730 -4.7738576 0.040328258 Taeniodus? 141 19740 0.074445618 -2.5976864 0.193386368 Venustodus 1 0 0.000527983 -7.5464463 0.003984396

Grand Total: 1,894 351,010 2.875062451

Table TA3. Density calculations by species class for the 213 Quarry.

Top No. of No. of No. of No. of Total No. of No. of 213 Quarry Community Slab indv. indv. Bottom indv. indv. (Bottom indv. indv. Density by Species Class (Top species species of Slab species species + Top species species Density: n / Area & per ft2 per m2 per ft2 per m2 Slabs) per ft2 per m2 Loose)

Anthozoa (1) Zaphrentoides spinulosus 6 0.17 1.85 1 0.03 0.31 7 0.20 2.15

Scyphozoa? (1) Sphenothallus sp. 4 0.11 1.23 4 0.11 1.23 8 0.23 2.46

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Pelecypoda (1) Sulcatopinna missouriensis 1 0.03 0.31 0 0.00 0.00 1 0.03 0.31

Polychaeta? (1) Crinicaminus haneyensis 1 0.03 0.31 0 0.00 0.00 1 0.03 0.31

Ostracoda (3) Amphissites? 1 0.03 0.31 0 0.00 0.00 1 0.03 0.31 Bairdia? 1 0.03 0.31 0 0.00 0.00 1 0.03 0.31 Coronakirkbya? 3 0.09 0.92 0 0.00 0.00 3 0.09 0.92

Articulata (6) Anthracospirifer leidyi 201 5.74 61.82 115 3.29 35.37 316 9.03 97.18 Cleiothyridina sublamellosa 39 1.11 11.99 16 0.46 4.92 55 1.57 16.91 Composita lewisensis 9 0.26 2.77 10 0.29 3.08 19 0.54 5.84

Table TA3. (Continued)

Composita subquadrata 40 1.14 12.30 25 0.71 7.69 65 1.86 19.99 Diaphragmus elegans 16 0.46 4.92 7 0.20 2.15 23 0.66 7.07 Eumetria verneuliana 3 0.09 0.92 0 0.00 0.00 3 0.09 0.92

Inarticulata (2) Oehlertella pleurites 2 0.06 0.62 0 0.00 0.00 2 0.06 0.62 Orbiculoidea sp. 6 0.17 1.85 2 0.06 0.62 8 0.23 2.46

Stenolaemata (9) Archimedes cf. meekanus 1525 43.57 469.00 115 3.29 35.37 1738 49.66 534.51 Eridopora sp. 11 0.31 3.38 16 0.46 4.92 16 0.46 4.92 Fenestella sp. 11 0.31 3.38 10 0.29 3.08 12 0.34 3.69

238 Glyptopora punctipora 111 3.17 34.14 25 0.71 7.69 135 3.86 41.52

Lyroporella sp. 1 0.03 0.31 7 0.20 2.15 1 0.03 0.31 Polypora sp. 85 2.43 26.14 0 0.00 0.00 99 2.83 30.45 Rhombopora sp. 261 7.46 80.27 0 0.00 0.00 432 12.34 132.86 Septopora sp. 3 0.09 0.92 2 0.06 0.62 8 0.23 2.46 Sulcoretepora sp. 3 0.09 0.92 0 0.00 0.00 4 0.11 1.23

Blastoidea (1) Pentremities elegans 29 0.83 8.92 12 0.34 3.69 41 1.17 12.61

Echinoidea (2) Lepidesthes formosa 1 0.03 0.31 0 0.00 0.00 1 0.03 0.31 Archaeocidaris megastylus 0 0.00 0.00 3 0.09 0.92 3 0.09 0.92 (isolated spines only)

Crinoidea (16)

Table TA3. (Continued)

Acrocrinus shumardi 1 0.03 0.31 0 0.00 0.00 1 0.03 0.31 Agassizocrinus lobatus 1 0.03 0.31 0 0.00 0.00 1 0.03 0.31 (juvenile) Anartiocrinus maxvillensis 1 0.03 0.31 0 0.00 0.00 1 0.03 0.31 Aphelecrinus mundus 5 0.14 1.54 0 0.00 0.00 5 0.14 1.54 Camptocrinus cirrifer 11 0.31 3.38 1 0.03 0.31 12 0.34 3.69 Cryphiocrinus girtyi 2 0.06 0.62 0 0.00 0.00 2 0.06 0.62 Cymbiocrinus grandis 54 1.54 16.61 1 0.03 0.31 55 1.57 16.91 Cymbiocrinus tumidus 11 0.31 3.38 0 0.00 0.00 11 0.31 3.38 Dasciocrinus florealis 30 0.86 9.23 5 0.14 1.54 35 1.00 10.76 Pentaramicrinus gracilis 25 0.71 7.69 3 0.09 0.92 28 0.80 8.61 Phanocrinus bimagnaramus 27 0.77 8.30 2 0.06 0.62 29 0.83 8.92 Phanocrinus maniformis 4 0.11 1.23 0 0.00 0.00 4 0.11 1.23

239 Pterotocrinus acutus 10 0.29 3.08 6 0.17 1.85 16 0.46 4.92

Taxocrinus whitfieldi 29 0.83 8.92 2 0.06 0.62 31 0.89 9.53 Tholocrinus spinosus 4 0.11 1.23 0 0.00 0.00 4 0.11 1.23 Zeacrinites magnoliaeformis 3 0.09 0.92 0 0.00 0.00 3 0.09 0.92

Chondrichthyes (2) Cladodus 1 0.03 0.31 0 0.00 0.00 1 0.03 0.31 Venustodus 1 0.03 0.31 0 0.00 0.00 1 0.03 0.31

Grand Total: 2,594 649 3,243

Table TA4. Density calculations for the Valley Stone Quarry.

Crs.- Total Fine-grained, Valley Stone Quarry No. of No. of Crs.- No. of No. of grained, No. of No. of No. of No. of No. of No. of (Top, calcarenites Arg. Community Density by indv. indv. grained, indv. indv. arg. indv. indv. indv. indv. indv. indv. Bottom and intbd calcilutites Species Class species species calcarenite species species calcarenite species species species species species species & calcilutite (Basinal) Density: n / Area per ft2 per m2 (Shoal) per ft2 per m2 (Shoal- per ft2 per m2 per ft2 per m2 per ft2 per m2 Loose) (Transitional) Margin)

Demospongea (1) Belemnospongia fascicularis 233 5.32 57.31 134 6.24 67.18 16 1.66 17.83 8 1.29 13.84 75 11.70 125.94

Anthozoa (1) Unidentifiable tabulate 1 0.02 0.25 1 0.05 0.50 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 coral?

240 Scyphozoa? (2)

Sphenothallus sp. 28 0.64 6.89 4 0.19 2.01 13 1.35 14.49 9 1.45 15.57 2 0.31 3.36 Paraconularia missouriensis 8 0.18 1.97 3 0.14 1.50 0 0.00 0.00 0 0.00 0.00 5 0.78 8.40

Pelecypoda (1) Sulcatopinna missouriensis 4 0.09 0.98 1 0.05 0.50 0 0.00 0.00 0 0.00 0.00 3 0.47 5.04

Polychaeta? (1) Crinicaminus haneyensis 2 0.05 0.49 2 0.09 1.00 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00

Articulata (11) Anthracospirifer leidyi 272 6.22 66.91 166 7.73 83.22 26 2.69 28.97 38 6.11 65.76 42 6.55 70.53 Cleiothyridina sublamellosa 8 0.18 1.97 0 0.00 0.00 2 0.21 2.23 3 0.48 5.19 3 0.47 5.04 Composita lewisensis 8 0.18 1.97 5 0.23 2.51 1 0.10 1.11 0 0.00 0.00 2 0.31 3.36 Composita subquadrata 410 9.37 100.85 63 2.93 31.58 215 22.26 239.57 25 4.02 43.26 107 16.69 179.68 Diaphragmus elegans 26 0.59 6.40 1 0.05 0.50 3 0.31 3.34 1 0.16 1.73 21 3.28 35.26 Dielasma sp. 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Eumetria costata 4 0.09 0.98 2 0.09 1.00 0 0.00 0.00 0 0.00 0.00 2 0.31 3.36 Eumetria verneuliana 5 0.11 1.23 2 0.09 1.00 1 0.10 1.11 0 0.00 0.00 2 0.31 3.36 Orthotetes kaskaskiensis 7 0.16 1.72 2 0.09 1.00 0 0.00 0.00 0 0.00 0.00 5 0.78 8.40

Table TA4. (Continued)

Punctospirifer sp. 7 0.16 1.72 0 0.00 0.00 2 0.21 2.23 0 0.00 0.00 5 0.78 8.40 Streptorhynchus sp. 1 0.02 0.25 1 0.05 0.50 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00

Inarticulata (2) Lingulipora sp. 2 0.05 0.49 2 0.09 1.00 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 Orbiculoidea sp. 6 0.14 1.48 1 0.05 0.50 2 0.21 2.23 3 0.48 5.19 0 0.00 0.00

Stenolaemata (10) Anisotrypa sp. 3 0.07 0.74 2 0.09 1.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Archimedes cf. meekanus 17 0.39 4.18 11 0.51 5.51 6 0.62 6.69 0 0.00 0.00 0 0.00 0.00 Eridopora sp. 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Fenestella sp. 4 0.09 0.98 0 0.00 0.00 0 0.00 0.00 1 0.16 1.73 3 0.47 5.04 Fenestrellina sp. 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 1 0.16 1.73 0 0.00 0.00 Polypora sp. 24 0.55 5.90 2 0.09 1.00 4 0.41 4.46 5 0.80 8.65 13 2.03 21.83 Rhombopora sp. 45 1.03 11.07 13 0.61 6.52 4 0.41 4.46 0 0.00 0.00 28 4.37 47.02 Septopora sp. 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68

241 Thamniscus? 40 0.91 9.84 16 0.75 8.02 7 0.72 7.80 14 2.25 24.23 3 0.47 5.04

Unidentifiable encrusting 6 0.14 1.48 5 0.23 2.51 1 0.10 1.11 0 0.00 0.00 0 0.00 0.00 bryozoan

Edrioasteroidea (2) Unidentifiable edrioasteroid 6 0.14 1.48 6 0.28 3.01 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 Lepidodiscus laudoni 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68

Blastoidea (1) Pentremities elegans 89 2.03 21.89 29 1.35 14.54 8 0.83 8.91 2 0.32 3.46 50 7.80 83.96

Echinoidea (1) Archaeocidaris megastylus 54 1.23 13.28 1 0.05 0.50 0 0.00 0.00 14 2.25 24.23 39 6.08 65.49

Ophiuroidea (1) Unidentifiable 34 0.78 8.36 21 0.98 10.53 2 0.21 2.23 9 1.45 15.57 2 0.31 3.36 encrinasterid brittle star

Crinoidea (18) Agassizocrinus lobatus 30 0.69 7.38 3 0.14 1.50 5 0.52 5.57 3 0.48 5.19 19 2.96 31.91

Table TA4. (Continued)

Anartiocrinus maxvillensis 31 0.71 7.63 21 0.98 10.53 0 0.00 0.00 3 0.48 5.19 7 1.09 11.75 Aphelecrinus mundus 25 0.57 6.15 14 0.65 7.02 7 0.72 7.80 0 0.00 0.00 4 0.62 6.72 Camptocrinus cirrifer 2 0.05 0.49 0 0.00 0.00 1 0.10 1.11 1 0.16 1.73 0 0.00 0.00 Cryphiocrinus girtyi 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Cymbiocrinus grandis 35 0.80 8.61 15 0.70 7.52 7 0.72 7.80 0 0.00 0.00 13 2.03 21.83 Cymbiocrinus tumidus 23 0.53 5.66 8 0.37 4.01 3 0.31 3.34 0 0.00 0.00 12 1.87 20.15 Dasciocrinus florealis 88 2.01 21.65 46 2.14 23.06 8 0.83 8.91 12 1.93 20.77 22 3.43 36.94 Eupachycrinus boydii 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Linocrinus wachsmuthi 48 1.10 11.81 36 1.68 18.05 4 0.41 4.46 5 0.80 8.65 3 0.47 5.04 Pentaramicrinus gracilis 16 0.37 3.94 2 0.09 1.00 0 0.00 0.00 0 0.00 0.00 14 2.18 23.51 Phanocrinus bimagnaramus 141 3.22 34.68 68 3.17 34.09 24 2.48 26.74 26 4.18 44.99 23 3.59 38.62 Phanocrinus maniformis 1 0.02 0.25 1 0.05 0.50 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 Phacelocrinus longidactylus 1 0.02 0.25 0 0.00 0.00 1 0.10 1.11 0 0.00 0.00 0 0.00 0.00 Platycrinites sp. 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Talarocrinus sp. 6 0.14 1.48 4 0.19 2.01 0 0.00 0.00 2 0.32 3.46 0 0.00 0.00 Taxocrinus whitfieldi 11 0.25 2.71 8 0.37 4.01 0 0.00 0.00 0 0.00 0.00 3 0.47 5.04

24 Tholocrinus spinosus 1 0.02 0.25 1 0.05 0.50 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00

Chondrichthyes (13) Agassizodus 1 0.02 0.25 0 0.00 0.00 1 0.10 1.11 0 0.00 0.00 0 0.00 0.00 Cladodus 3 0.07 0.74 0 0.00 0.00 1 0.10 1.11 0 0.00 0.00 2 0.31 3.36 Ctenacanthus 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Deltodopsis 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Deltodus 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Listracanthus? 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Lophodus 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Mesodmodus 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Petrodus 59 1.35 14.51 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 59 9.20 99.08 Psammodus? 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Polyrhizodus 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Taeniodus? 1 0.02 0.25 0 0.00 0.00 0 0.00 0.00 0 0.00 0.00 1 0.16 1.68 Venustodus 1 0.02 0.25 0 0.00 0.00 1 0.10 1.11 0 0.00 0.00 0 0.00 0.00

Grand Total: 1,894 723 376 185 610

Table TA5. Simpson’s Index of Diversity and Shannon’s Evenness calculations for the 213 Quarry, top-of-slab.

Proportion of entire Natural Log Number of population of 213 Quarry, Top-of-slab - Individuals to Proportion Simpson's Index of Diversity individual and Shannon's Evenness species Calculations Top-of-slab Top-of- Top-of-slab Top-of-slab Top-of-slab (nT) (top + slab (PT*ln PT *- (PT) (ln PT) loose total) nT(nT - 1) 1)

Anthozoa (1)

243 Zaphrentoides spinulosus 6 30 0.002313030 -6.069196896 0.014038235

Scyphozoa? (1) Sphenothallus sp. 4 12 0.001542020 -6.474662004 0.009984059

Pelecypoda (1) Sulcatopinna missouriensis 1 0 0.000385505 -7.860956365 0.003030438

Polychaeta? (1) Crinicaminus haneyensis 1 0 0.000385505 -7.860956365 0.003030438

Ostracoda (3) Amphissites? 1 0 0.000385505 -7.860956365 0.003030438 Bairdia? 1 0 0.000385505 -7.860956365 0.003030438 Coronakirkbya? 3 6 0.001156515 -6.762344076 0.007820753

Table TA5. (Continued)

Articulata (6) Anthracospirifer leidyi 201 40200 0.077486507 -2.557651457 0.198183478 Cleiothyridina sublamellosa 39 1482 0.015034695 -4.197394719 0.063106551 Composita lewisensis 9 72 0.003469545 -5.663731788 0.019650573 Composita subquadrata 40 1560 0.015420200 -4.172076911 0.064334262 Diaphragmus elegans 16 240 0.006168080 -5.088367643 0.031385460 Eumetria verneuliana 3 6 0.001156515 -6.762344076 0.007820753

Inarticulata (2) Oehlertella pleurites 2 2 0.000771010 -7.167809184 0.005526453 Orbiculoidea sp. 6 30 0.002313030 -6.069196896 0.014038235

244 Stenolaemata (9)

Archimedes cf. meekanus 1525 2324100 0.587895143 -0.531206676 0.312293824 Eridopora sp. 11 110 0.004240555 -5.463061092 0.023166412 Fenestella sp. 11 110 0.004240555 -5.463061092 0.023166412 Glyptopora punctipora 111 12210 0.042791056 -3.151426164 0.134852854 Lyroporella sp. 1 0 0.000385505 -7.860956365 0.003030438 Polypora sp. 85 7140 0.032767926 -3.418305108 0.112010769 Rhombopora sp. 261 67860 0.100616808 -2.296435958 0.231060056 Septopora sp. 3 6 0.001156515 -6.762344076 0.007820753 Sulcoretepora sp. 3 6 0.001156515 -6.762344076 0.007820753

Blastoidea (1) Pentremities elegans 29 812 0.011179645 -4.493660535 0.050237531

Echinoidea (2) Lepidesthes formosa 1 0 0.000385505 -7.860956365 0.003030438

Table TA5. (Continued) ln of a Archaeocidaris megastylus number ≤ 0 is (isolated spines only) 0 0 0 undefined

Crinoidea (16) Acrocrinus shumardi 1 0 0.000385505 -7.860956365 0.003030438 Agassizocrinus lobatus (juvenile) 1 0 0.000385505 -7.860956365 0.003030438 Anartiocrinus maxvillensis 1 0 0.000385505 -7.860956365 0.003030438 Aphelecrinus mundus 5 20 0.001927525 -6.251518452 0.012049958 Camptocrinus cirrifer 11 110 0.004240555 -5.463061092 0.023166412 Cryphiocrinus girtyi 2 2 0.000771010 -7.167809184 0.005526453 Cymbiocrinus grandis 54 2862 0.020817271 -3.871972318 0.080603896 245 Cymbiocrinus tumidus 11 110 0.004240555 -5.463061092 0.023166412

Dasciocrinus florealis 30 870 0.011565150 -4.459758983 0.051577783 Pentaramicrinus gracilis 25 600 0.009637625 -4.642080540 0.044738633 Phanocrinus bimagnaramus 27 702 0.010408635 -4.565119499 0.047516664 Phanocrinus maniformis 4 12 0.001542020 -6.474662004 0.009984059 Pterotocrinus acutus 10 90 0.003855050 -5.558371272 0.021427800 Taxocrinus whitfieldi 29 812 0.011179645 -4.493660535 0.050237531 Tholocrinus spinosus 4 12 0.001542020 -6.474662004 0.009984059 Zeacrinites magnoliaeformis 3 6 0.001156515 -6.762344076 0.007820753

Chondrichthyes (2) Cladodus 1 0 0.000385505 -7.860956365 0.003030438 Venustodus 1 0 0.000385505 -7.860956365 0.003030438

Grand Total of N 2,594 2,462,202 1.759453404

Table TA6. Simpson’s Index of Diversity and Shannon’s Evenness calculations for the 213 Quarry, bottom-of-slab.

Proportion of entire Number of population Natural Log of Proportion 213 Quarry, Bottom-of-slab - Individuals to Simpson's Index of Diversity individual and Shannon's Evenness species Calculations Bottom- Bottom-of- Bottom-of- Bottom-of- of-slab Bottom-of-slab (ln PB) slab (PB * ln slab (nB) slab (PB) nB(nB - 1) PB * -1)

Anthozoa (1)

246 Zaphrentoides spinulosus 1 0 0.001540832 -6.475432717 0.009977554

Scyphozoa? (1) Sphenothallus sp. 4 12 0.006163328 -5.089138356 0.031366030

Pelecypoda (1) ln of a number ≤ 0 is Sulcatopinna missouriensis 0 0 0 undefined

Polychaeta? (1) ln of a number ≤ 0 is Crinicaminus haneyensis 0 0 0 undefined

Ostracoda (3)

Table TA6. (Continued) ln of a number ≤ 0 is Amphissites? 0 0 0 undefined ln of a number ≤ 0 is Bairdia? 0 0 0 undefined ln of a number ≤ 0 is Coronakirkbya? 0 0 0 undefined

Articulata (6) Anthracospirifer leidyi 115 13110 0.177195686 -1.730500588 0.306637238 Cleiothyridina sublamellosa 16 240 0.024653313 -3.702843994 0.091287371 Composita lewisensis 10 90 0.015408320 -4.172847624 0.064296574 Composita subquadrata 25 600 0.038520801 -3.256556892 0.125445181 Diaphragmus elegans 7 42 0.010785824 -4.529522568 0.048854635

247 ln of a number ≤ 0 is

Eumetria verneuliana 0 0 0 undefined

Inarticulata (2) ln of a number ≤ 0 is Oehlertella pleurites 0 0 0 undefined Orbiculoidea sp. 2 2 0.003081664 -5.782285536 0.017819062

Stenolaemata (9) Archimedes cf. meekanus 213 45156 0.328197227 -1.114140551 0.365657839 Eridopora sp. 5 20 0.007704160 -4.865994804 0.037488404 Fenestella sp. 1 0 0.001540832 -6.475432717 0.009977554 Glyptopora punctipora 24 552 0.036979969 -3.297378886 0.121936970 ln of a number ≤ 0 is Lyroporella sp. 0 0 0 undefined Polypora sp. 14 182 0.021571649 -3.836375387 0.082756942

Table TA6. (Continued)

Rhombopora sp. 171 29070 0.263482280 -1.333769160 0.351424540 Septopora sp. 5 20 0.007704160 -4.865994804 0.037488404 Sulcoretepora sp. 1 0 0.001540832 -6.475432717 0.009977554

Blastoidea (1) Pentremities elegans 12 132 0.018489985 -3.990526067 0.073784765

Echinoidea (2) ln of a number ≤ 0 is Lepidesthes formosa 0 0 0 undefined Archaeocidaris megastylus (isolated spines only) 3 6 0.004622496 -5.376820428 0.024854332

248 Crinoidea (16) ln of a number ≤ 0 is Acrocrinus shumardi 0 0 0 undefined Agassizocrinus lobatus ln of a number ≤ 0 is (juvenile) 0 0 0 undefined ln of a number ≤ 0 is Anartiocrinus maxvillensis 0 0 0 undefined ln of a number ≤ 0 is Aphelecrinus mundus 0 0 0 undefined Camptocrinus cirrifer 1 0 0.001540832 -6.475432717 0.009977554 ln of a number ≤ 0 is Cryphiocrinus girtyi 0 0 0 undefined Cymbiocrinus grandis 1 0 0.001540832 -6.475432717 0.009977554 ln of a number ≤ 0 is Cymbiocrinus tumidus 0 0 0 undefined Dasciocrinus florealis 5 20 0.007704160 -4.865994804 0.037488404

Table TA6. (Continued)

Pentaramicrinus gracilis 3 6 0.004622496 -5.376820428 0.024854332 Phanocrinus bimagnaramus 2 2 0.003081664 -5.782285536 0.017819062 ln of a number ≤ 0 is Phanocrinus maniformis 0 0 0 undefined Pterotocrinus acutus 6 30 0.009244992 -4.683673247 0.043300523 Taxocrinus whitfieldi 2 2 0.003081664 -5.782285536 0.017819062 ln of a number ≤ 0 is Tholocrinus spinosus 0 0 0 undefined ln of a number ≤ 0 is Zeacrinites magnoliaeformis 0 0 0 undefined

Chondrichthyes (2) ln of a number ≤ 0 is 249 Cladodus 0 0 0 undefined

ln of a number ≤ 0 is Venustodus 0 0 0 undefined

Grand Total of N 649 89,294 1.972267439

Table TA7. Simpson’s Index of Diversity and Shannon’s Evenness calculations for the Valley Stone Quarry, coarse-grained calcarenite (shoal).

Proportion of entire Valley Stone Quarry, Coarse- Number of population Natural Log of Proportion grained calcarenite (Shoal) - Individuals to Simpson's Index of Diversity and individual Shannon's Evenness Calculations species Shoal Shoal (P S * Shoal nS Shoal (PS) Shoal (ln PS) nST(nS - 1) ln P S * -1)

Demospongea (1) Belemnospongia fascicularis 134 17822 0.185338866 -1.685569422 0.312401525

250

Anthozoa (1) Unidentifiable tabulate coral? 1 0 0.001383126 -6.583409222 0.009105684

Scyphozoa? (2) Sphenothallus sp. 4 12 0.005532503 -5.197114861 0.028753056 Paraconularia missouriensis 3 6 0.004149378 -5.484796933 0.022758493

Pelecypoda (1) Sulcatopinna missouriensis 1 0 0.001383126 -6.583409222 0.009105684

Polychaeta? (1) Crinicaminus haneyensis 2 2 0.002766252 -5.890262042 0.016293948

Articulata (11)

Table TA7. (Continued)

Anthracospirifer leidyi 166 27390 0.229598893 -1.471421434 0.337836733 ln of a number ≤ 0 is 0 0 0 Cleiothyridina sublamellosa undefined Composita lewisensis 5 20 0.006915629 -4.973971310 0.034398142 Composita subquadrata 63 3906 0.087136929 -2.440274496 0.212638027 Diaphragmus elegans 1 0 0.001383126 -6.583409222 0.009105684 ln of a number ≤ 0 is 0 0 0 Dielasma sp. undefined Eumetria costata 2 2 0.002766252 -5.890262042 0.016293948 Eumetria verneuliana 2 2 0.002766252 -5.890262042 0.016293948 Orthotetes kaskaskiensis 2 2 0.002766252 -5.890262042 0.016293948 ln of a number ≤ 0 is 0 0 0 Punctospirifer sp. undefined

251 Streptorhynchus sp. 1 0 0.001383126 -6.583409222 0.009105684

Inarticulata (2) Lingulipora sp. 2 2 0.002766252 -5.890262042 0.016293948 Orbiculoidea sp. 1 0 0.001383126 -6.583409222 0.009105684

Stenolaemata (10) Anisotrypa sp. 2 2 0.002766252 -5.890262042 0.016293948 Archimedes cf. meekanus 11 110 0.015214385 -4.185513949 0.063680019 ln of a number ≤ 0 is 0 0 0 Eridopora sp. undefined ln of a number ≤ 0 is 0 0 0 Fenestella sp. undefined ln of a number ≤ 0 is 0 0 0 Fenestrellina sp. undefined Polypora sp. 2 2 0.002766252 -5.890262042 0.016293948

Table TA7. (Continued)

Rhombopora sp. 13 156 0.017980636 -4.018459865 0.072254465 ln of a number ≤ 0 is 0 0 0 Septopora sp. undefined Thamniscus? 16 240 0.022130014 -3.810820500 0.084333510 Unidentifiable encrusting bryozoan 5 20 0.006915629 -4.973971310 0.034398142

Edrioasteroidea (2) Unidentifiable edrioasteroid 6 30 0.008298755 -4.791649753 0.039764728 ln of a number ≤ 0 is 0 0 0 Lepidodiscus laudoni undefined

Blastoidea (1) Pentremities elegans 29 812 0.040110650 -3.216113392 0.129000399 252 Echinoidea (1) Archaeocidaris megastylus 1 0 0.001383126 -6.583409222 0.009105684

Ophiuroidea (1) Unidentifiable encrinasterid brittle 21 420 0.029045643 -3.538886784 0.102789243 star

Crinoidea (18) Agassizocrinus lobatus 3 6 0.004149378 -5.484796933 0.022758493 Anartiocrinus maxvillensis 21 420 0.029045643 -3.538886784 0.102789243 Aphelecrinus mundus 14 182 0.019363762 -3.944351893 0.076377492 ln of a number ≤ 0 is 0 0 0 Camptocrinus cirrifer undefined ln of a number ≤ 0 is 0 0 0 Cryphiocrinus girtyi undefined

Table TA7. (Continued)

Cymbiocrinus grandis 15 210 0.020746888 -3.875359021 0.080401639 Cymbiocrinus tumidus 8 56 0.011065007 -4.503967680 0.049836434 Dasciocrinus florealis 46 2070 0.063623790 -2.754767826 0.175268769 ln of a number ≤ 0 is 0 0 0 Eupachycrinus boydii undefined Linocrinus wachsmuthi 36 1260 0.049792531 -2.999890284 0.149372130 Pentaramicrinus gracilis 2 2 0.002766252 -5.890262042 0.016293948 Phanocrinus bimagnaramus 68 4556 0.094052559 -2.363901517 0.222330986 Phanocrinus maniformis 1 0 0.001383126 -6.583409222 0.009105684 ln of a number ≤ 0 is 0 0 0 Phacelocrinus longidactylus undefined ln of a number ≤ 0 is 0 0 0 Platycrinites sp. undefined

253 Talarocrinus sp. 4 12 0.005532503 -5.197114861 0.028753056

Taxocrinus whitfieldi 8 56 0.011065007 -4.503967680 0.049836434 Tholocrinus spinosus 1 0 0.001383126 -6.583409222 0.009105684

Table TA7. (Continued) ln of a number ≤ 0 is 0 0 0 Lophodus undefined ln of a number ≤ 0 is 0 0 0 Mesodmodus undefined ln of a number ≤ 0 is 0 0 0 Petrodus undefined ln of a number ≤ 0 is 0 0 0 Psammodus? undefined ln of a number ≤ 0 is 0 0 0 Polyrhizodus undefined ln of a number ≤ 0 is 0 0 0 Taeniodus? undefined ln of a number ≤ 0 is 0 0 0 Venustodus undefined

254

Grand Total of N: 723 59,788 2.635928207

Table TA8. Simpson’s Index of Diversity and Shannon’s Evenness calculations for the Valley Stone Quarry, coarse-grained, argillaceous calcarenites (Shoal-margin).

Proportion of entire Number of population Valley Stone Quarry, Coarse- Natural Log of Proportion Individuals to grained, argillaceous calcarenites individual (Shoal-Margin) - Simpson's Index species of Diversity and Shannon's Shoal- Evenness Calculations Shoal- Shoal- Shoal- Margin (P SM Margin Margin Margin Shoal-Margin (ln PSM) * ln P SM * - nSM nSM(nSM - 1) (PSM) 1)

255 Demospongea (1)

Belemnospongia fascicularis 16 240 0.042553191 -3.157000421 0.134340443

Anthozoa (1) ln of a number ≤ 0 is 0 0 0 Unidentifiable tabulate coral? undefined

Scyphozoa? (2) Sphenothallus sp. 13 156 0.034574468 -3.364639786 0.116330631 ln of a number ≤ 0 is 0 0 0 Paraconularia missouriensis undefined

Table TA8. (Continued)

Pelecypoda (1) ln of a number ≤ 0 is 0 0 0 Sulcatopinna missouriensis undefined

Polychaeta? (1) ln of a number ≤ 0 is 0 0 0 Crinicaminus haneyensis undefined

Articulata (11) Anthracospirifer leidyi 26 650 0.069148936 -2.671492605 0.184730872 Cleiothyridina sublamellosa 2 2 0.005319149 -5.236441963 0.027853415 Composita lewisensis 1 0 0.002659574 -5.929589143 0.015770184 Composita subquadrata 215 46010 0.571808511 -0.558951115 0.319613005 256 Diaphragmus elegans 3 6 0.007978723 -4.830976855 0.038545028 ln of a number ≤ 0 is 0 0 0 Dielasma sp. undefined ln of a number ≤ 0 is 0 0 0 Eumetria costata undefined Eumetria verneuliana 1 0 0.002659574 -5.929589143 0.015770184 ln of a number ≤ 0 is 0 0 0 Orthotetes kaskaskiensis undefined Punctospirifer sp. 2 2 0.005319149 -5.236441963 0.027853415 ln of a number ≤ 0 is 0 0 0 Streptorhynchus sp. undefined

Inarticulata (2) ln of a number ≤ 0 is 0 0 0 Lingulipora sp. undefined Orbiculoidea sp. 2 2 0.005319149 -5.236441963 0.027853415

Table TA8. (Continued)

Stenolaemata (10) ln of a number ≤ 0 is 0 0 0 Anisotrypa sp. undefined Archimedes cf. meekanus 6 30 0.015957447 -4.137829674 0.066029197 ln of a number ≤ 0 is 0 0 0 Eridopora sp. undefined ln of a number ≤ 0 is 0 0 0 Fenestella sp. undefined ln of a number ≤ 0 is 0 0 0 Fenestrellina sp. undefined Polypora sp. 4 12 0.010638298 -4.543294782 0.048332923 Rhombopora sp. 4 12 0.010638298 -4.543294782 0.048332923 ln of a number ≤ 0 is 0 0 0 Septopora sp. undefined

257 Thamniscus? 7 42 0.018617021 -3.983678994 0.074164237

Unidentifiable encrusting bryozoan 1 0 0.002659574 -5.929589143 0.015770184

Edrioasteroidea (2) ln of a number ≤ 0 is 0 0 0 Unidentifiable edrioasteroid undefined ln of a number ≤ 0 is 0 0 0 Lepidodiscus laudoni undefined

Blastoidea (1) Pentremities elegans 8 56 0.021276596 -3.850147602 0.081918034

Echinoidea (1) ln of a number ≤ 0 is 0 0 0 Archaeocidaris megastylus undefined

Table TA8. (Continued)

Ophiuroidea (1) Unidentifiable encrinasterid brittle 2 2 0.005319149 -5.236441963 0.027853415 star

Crinoidea (18) Agassizocrinus lobatus 5 20 0.013297872 -4.320151231 0.057448820 ln of a number ≤ 0 is 0 0 0 Anartiocrinus maxvillensis undefined Aphelecrinus mundus 7 42 0.018617021 -3.983678994 0.074164237 Camptocrinus cirrifer 1 0 0.002659574 -5.929589143 0.015770184 ln of a number ≤ 0 is 0 0 0 Cryphiocrinus girtyi undefined Cymbiocrinus grandis 7 42 0.018617021 -3.983678994 0.074164237

258 Cymbiocrinus tumidus 3 6 0.007978723 -4.830976855 0.038545028

Dasciocrinus florealis 8 56 0.021276596 -3.850147602 0.081918034 ln of a number ≤ 0 is 0 0 0 Eupachycrinus boydii undefined Linocrinus wachsmuthi 4 12 0.010638298 -4.543294782 0.048332923 ln of a number ≤ 0 is 0 0 0 Pentaramicrinus gracilis undefined Phanocrinus bimagnaramus 24 552 0.063829787 -2.751535313 0.175629914 ln of a number ≤ 0 is 0 0 0 Phanocrinus maniformis undefined Phacelocrinus longidactylus 1 0 0.002659574 -5.929589143 0.015770184 ln of a number ≤ 0 is 0 0 0 Platycrinites sp. undefined ln of a number ≤ 0 is 0 0 0 Talarocrinus sp. undefined ln of a number ≤ 0 is 0 0 0 Taxocrinus whitfieldi undefined

Table TA8. (Continued) ln of a number ≤ 0 is 0 0 0 Tholocrinus spinosus undefined

Chondrichthyes (13) Agassizodus 1 0 0.002659574 -5.929589143 0.015770184 Cladodus 1 0 0.002659574 -5.929589143 0.015770184 ln of a number ≤ 0 is 0 0 0 Ctenacanthus undefined ln of a number ≤ 0 is 0 0 0 Deltodopsis undefined ln of a number ≤ 0 is 0 0 0 Deltodus undefined ln of a number ≤ 0 is 0 0 0 Listracanthus? undefined

259 ln of a number ≤ 0 is 0 0 0 Lophodus undefined ln of a number ≤ 0 is 0 0 0 Mesodmodus undefined ln of a number ≤ 0 is 0 0 0 Petrodus undefined ln of a number ≤ 0 is 0 0 0 Psammodus? undefined ln of a number ≤ 0 is 0 0 0 Polyrhizodus undefined ln of a number ≤ 0 is 0 0 0 Taeniodus? undefined Venustodus 1 0 0.002659574 -5.929589143 0.015770184

Grand Total of N: 376 47,952 1.884345431

Table TA9. Simpson’s Index of Diversity and Shannon’s Evenness calculations for the Valley Stone Quarry, fine-grained calcarenites and interbedded calcilutites (Transitional).

Proportion of entire Number of population Natural Log of Proportion Valley Stone Quarry, Fine- Individuals to grained calcarenites and individual interbedded calcilutites species (Transitional) - Simpson's Index of Diversity and Shannon's Transitional Evenness Calculations Transitional Transitional Transitional Transitional (ln PT) (P T * ln P T * nT nT(nT - 1) (PT) -1)

260 Demospongea (1) Belemnospongia fascicularis 8 56 0.043243243 -3.140914283 0.13582332

Anthozoa (1) ln of a number ≤ 0 is 0 0 0 Unidentifiable tabulate coral? undefined

Scyphozoa? (2) Sphenothallus sp. 9 72 0.048648649 -3.023131248 0.14707125 ln of a number ≤ 0 is 0 0 0 Paraconularia missouriensis undefined

Pelecypoda (1) ln of a number ≤ 0 is 0 0 0 Sulcatopinna missouriensis undefined

Table TA9. (Continued)

Polychaeta? (1) ln of a number ≤ 0 is 0 0 0 Crinicaminus haneyensis undefined

Articulata (11) Anthracospirifer leidyi 38 1406 0.205405405 -1.582769665 0.325109445 Cleiothyridina sublamellosa 3 6 0.016216216 -4.121743536 0.066839084 ln of a number ≤ 0 is 0 0 0 Composita lewisensis undefined Composita subquadrata 25 600 0.135135135 -2.001480000 0.270470270 Diaphragmus elegans 1 0 0.005405405 -5.220355825 0.028218140 ln of a number ≤ 0 is 0 0 0 Dielasma sp. undefined

261 ln of a number ≤ 0 is 0 0 0

Eumetria costata undefined ln of a number ≤ 0 is 0 0 0 Eumetria verneuliana undefined ln of a number ≤ 0 is 0 0 0 Orthotetes kaskaskiensis undefined ln of a number ≤ 0 is 0 0 0 Punctospirifer sp. undefined ln of a number ≤ 0 is 0 0 0 Streptorhynchus sp. undefined

Inarticulata (2) ln of a number ≤ 0 is 0 0 0 Lingulipora sp. undefined Orbiculoidea sp. 3 6 0.016216216 -4.121743536 0.066839084

Table TA9. (Continued)

Stenolaemata (10) ln of a number ≤ 0 is 0 0 0 Anisotrypa sp. undefined ln of a number ≤ 0 is 0 0 0 Archimedes cf. meekanus undefined ln of a number ≤ 0 is 0 0 0 Eridopora sp. undefined Fenestella sp. 1 0 0.005405405 -5.220355825 0.028218140 Fenestrellina sp. 1 0 0.005405405 -5.220355825 0.028218140 Polypora sp. 5 20 0.027027027 -3.610917913 0.097592376 ln of a number ≤ 0 is 0 0 0 Rhombopora sp. undefined ln of a number ≤ 0 is 0 0 0 Septopora sp. undefined

262 Thamniscus? 14 182 0.075675676 -2.581298495 0.195341508 ln of a number ≤ 0 is 0 0 0 Unidentifiable encrusting bryozoan undefined

Edrioasteroidea (2) ln of a number ≤ 0 is 0 0 0 Unidentifiable edrioasteroid undefined ln of a number ≤ 0 is 0 0 0 Lepidodiscus laudoni undefined

Blastoidea (1) Pentremities elegans 2 2 0.010810811 -4.527208645 0.048942796

Echinoidea (1) Archaeocidaris megastylus 14 182 0.075675676 -2.581298495 0.195341508

Table TA9. (Continued)

Ophiuroidea (1) Unidentifiable encrinasterid brittle 9 72 0.048648649 -3.023131248 0.14707125 star

Crinoidea (18) Agassizocrinus lobatus 3 6 0.016216216 -4.121743536 0.066839084 Anartiocrinus maxvillensis 3 6 0.016216216 -4.121743536 0.066839084 ln of a number ≤ 0 is 0 0 0 Aphelecrinus mundus undefined Camptocrinus cirrifer 1 0 0.005405405 -5.220355825 0.028218140 ln of a number ≤ 0 is 0 0 0 Cryphiocrinus girtyi undefined ln of a number ≤ 0 is 0 0 0

263 Cymbiocrinus grandis undefined ln of a number ≤ 0 is 0 0 0 Cymbiocrinus tumidus undefined Dasciocrinus florealis 12 132 0.064864865 -2.735449175 0.177434541 ln of a number ≤ 0 is 0 0 0 Eupachycrinus boydii undefined Linocrinus wachsmuthi 5 20 0.027027027 -3.610917913 0.097592376 ln of a number ≤ 0 is 0 0 0 Pentaramicrinus gracilis undefined Phanocrinus bimagnaramus 26 650 0.140540541 -1.962259287 0.275776981 ln of a number ≤ 0 is 0 0 0 Phanocrinus maniformis undefined ln of a number ≤ 0 is 0 0 0 Phacelocrinus longidactylus undefined ln of a number ≤ 0 is 0 0 0 Platycrinites sp. undefined Talarocrinus sp. 2 2 0.010810811 -4.527208645 0.048942796

Table TA9. (Continued) ln of a number ≤ 0 is 0 0 0 Taxocrinus whitfieldi undefined ln of a number ≤ 0 is 0 0 0 Tholocrinus spinosus undefined

Chondrichthyes (13) ln of a number ≤ 0 is 0 0 0 Agassizodus undefined ln of a number ≤ 0 is 0 0 0 Cladodus undefined ln of a number ≤ 0 is 0 0 0 Ctenacanthus undefined ln of a number ≤ 0 is 0 0 0 Deltodopsis undefined 264 ln of a number ≤ 0 is 0 0 0 Deltodus undefined ln of a number ≤ 0 is 0 0 0 Listracanthus? undefined ln of a number ≤ 0 is 0 0 0 Lophodus undefined ln of a number ≤ 0 is 0 0 0 Mesodmodus undefined ln of a number ≤ 0 is 0 0 0 Petrodus undefined ln of a number ≤ 0 is 0 0 0 Psammodus? undefined ln of a number ≤ 0 is 0 0 0 Polyrhizodus undefined ln of a number ≤ 0 is 0 0 0 Taeniodus? undefined

Table TA9. (Continued) ln of a number ≤ 0 is 0 0 0 Venustodus undefined

Grand Total of N: 185 3,420 2.542739313

Table TA10. Simpson’s Index of Diversity and Shannon’s Evenness calculations for the Valley Stone Quarry, argillaceous calcilutites (Basinal).

Proportion

265 of entire Number of population Natural Log of Proportion Valley Stone Quarry, Argillaceous Individuals to calcilutites (Basinal) - Simpson's individual Index of Diversity and Shannon's species Evenness Calculations Basinal Basinal (P Basinal nB nB(nB - Basinal (PB) Basinal (ln PB) B * ln P B * 1) -1)

Demospongea (1) Belemnospongia fascicularis 75 5550 0.12295082 -2.095970844 0.257701333

Anthozoa (1) ln of a number ≤ 0 is 0 0 0 Unidentifiable tabulate coral? undefined

Table TA10. (Continued)

Scyphozoa? (2) Sphenothallus sp. 2 2 0.003278689 -5.720311777 0.018755121 Paraconularia missouriensis 5 20 0.008196721 -4.804021045 0.039377222

Pelecypoda (1) Sulcatopinna missouriensis 3 6 0.004918033 -5.314846668 0.026138590

Polychaeta? (1) ln of a number ≤ 0 is 0 0 0 Crinicaminus haneyensis undefined

Articulata (11) Anthracospirifer leidyi 42 1722 0.068852459 -2.675789339 0.184234676

266 Cleiothyridina sublamellosa 3 6 0.004918033 -5.314846668 0.026138590

Composita lewisensis 2 2 0.003278689 -5.720311777 0.018755121 Composita subquadrata 107 11342 0.175409836 -1.740630123 0.305323644 Diaphragmus elegans 21 420 0.034426230 -3.368936519 0.115979782 Dielasma sp. 1 0 0.001639344 -6.413458957 0.010513867 Eumetria costata 2 2 0.003278689 -5.720311777 0.018755121 Eumetria verneuliana 2 2 0.003278689 -5.720311777 0.018755121 Orthotetes kaskaskiensis 5 20 0.008196721 -4.804021045 0.039377222 Punctospirifer sp. 5 20 0.008196721 -4.804021045 0.039377222 ln of a number ≤ 0 is 0 0 0 Streptorhynchus sp. undefined

Inarticulata (2) ln of a number ≤ 0 is 0 0 0 Lingulipora sp. undefined

Table TA10. (Continued) ln of a number ≤ 0 is 0 0 0 Orbiculoidea sp. undefined

Stenolaemata (10) Anisotrypa sp. 1 0 0.001639344 -6.413458957 0.010513867 ln of a number ≤ 0 is 0 0 0 Archimedes cf. meekanus undefined Eridopora sp. 1 0 0.001639344 -6.413458957 0.010513867 Fenestella sp. 3 6 0.004918033 -5.314846668 0.026138590 ln of a number ≤ 0 is 0 0 0 Fenestrellina sp. undefined Polypora sp. 13 156 0.021311475 -3.848509600 0.082017418 Rhombopora sp. 28 756 0.045901639 -3.081254447 0.141434630

267 Septopora sp. 1 0 0.001639344 -6.413458957 0.010513867

Thamniscus? 3 6 0.004918033 -5.314846668 0.026138590 ln of a number ≤ 0 is 0 0 0 Unidentifiable encrusting bryozoan undefined

Edrioasteroidea (2) ln of a number ≤ 0 is 0 0 0 Unidentifiable edrioasteroid undefined Lepidodiscus laudoni 1 0 0.001639344 -6.413458957 0.010513867

Blastoidea (1) Pentremities elegans 50 2450 0.081967213 -2.501435952 0.205035734

Echinoidea (1) Archaeocidaris megastylus 39 1482 0.063934426 -2.749897311 0.175813107

Table TA10. (Continued)

Ophiuroidea (1) Unidentifiable encrinasterid brittle 2 2 0.003278689 -5.720311777 0.018755121 star

Crinoidea (18) Agassizocrinus lobatus 19 342 0.031147541 -3.469019978 0.108051442 Anartiocrinus maxvillensis 7 42 0.011475410 -4.467548808 0.051266954 Aphelecrinus mundus 4 12 0.006557377 -5.027164596 0.032965014 ln of a number ≤ 0 is 0 0 0 Camptocrinus cirrifer undefined Cryphiocrinus girtyi 1 0 0.001639344 -6.413458957 0.010513867 Cymbiocrinus grandis 13 156 0.021311475 -3.848509600 0.082017418 Cymbiocrinus tumidus 12 132 0.019672131 -3.928552307 0.077282996 268 Dasciocrinus florealis 22 462 0.036065574 -3.322416504 0.119824858 Eupachycrinus boydii 1 0 0.001639344 -6.413458957 0.010513867 Linocrinus wachsmuthi 3 6 0.004918033 -5.314846668 0.026138590 Pentaramicrinus gracilis 14 182 0.022950820 -3.774401628 0.086625611 Phanocrinus bimagnaramus 23 506 0.037704918 -3.277964741 0.123595392 ln of a number ≤ 0 is 0 0 0 Phanocrinus maniformis undefined ln of a number ≤ 0 is 0 0 0 Phacelocrinus longidactylus undefined Platycrinites sp. 1 0 0.001639344 -6.413458957 0.010513867 ln of a number ≤ 0 is 0 0 0 Talarocrinus sp. undefined Taxocrinus whitfieldi 3 6 0.004918033 -5.314846668 0.026138590 ln of a number ≤ 0 is 0 0 0 Tholocrinus spinosus undefined

Table TA10. (Continued)

Chondrichthyes (13) ln of a number ≤ 0 is 0 0 0 Agassizodus undefined Cladodus 2 2 0.003278689 -5.720311777 0.018755121 Ctenacanthus 1 0 0.001639344 -6.413458957 0.010513867 Deltodopsis 1 0 0.001639344 -6.413458957 0.010513867 Deltodus 1 0 0.001639344 -6.413458957 0.010513867 Listracanthus? 1 0 0.001639344 -6.413458957 0.010513867 Lophodus 1 0 0.001639344 -6.413458957 0.010513867 Mesodmodus 1 0 0.001639344 -6.413458957 0.010513867 Petrodus 59 3422 0.096721311 -2.335921513 0.225933392 Psammodus? 1 0 0.001639344 -6.413458957 0.010513867 Polyrhizodus 1 0 0.001639344 -6.413458957 0.010513867

269 Taeniodus? 1 0 0.001639344 -6.413458957 0.010513867 ln of a number ≤ 0 is 0 0 0 Venustodus undefined

Grand Total of N: 610 29,242 2.941333071

Table TA11. Relative frequency calculations for the top-of-slab, bottom-of-slab, and entire landscape for the 213 Quarry.

213 213 Entire 213 Quarry, Relative 213 Top- Top-of-slab Bottom- Bottom- Landscape Frequency for Top-of-slab, Number of-slab (nT) (top + of-slab of-slab Relative Bottom-of-slab, and Entire (n) Relative loose total) (nB) Relative Frequency Frequency Landscape Frequency Total

Anthozoa (1) Zaphrentoides spinulosus 6 1 7 0.23% 0.15% 0.22%

Scyphozoa? (1) Sphenothallus sp. 4 4 8 0.15% 0.62% 0.22%

270 Pelecypoda (1)

Sulcatopinna missouriensis 1 0 1 0.04% 0.00% 0.03%

Polychaeta? (1) Crinicaminus haneyensis 1 0 1 0.04% 0.00% 0.03%

Ostracoda (3) Amphissites? 1 0 1 0.04% 0.00% 0.03% Bairdia? 1 0 1 0.04% 0.00% 0.03% Coronakirkbya? 3 0 3 0.12% 0.00% 0.09%

Articulata (6) Anthracospirifer leidyi 201 115 316 7.75% 17.72% 9.74% Cleiothyridina sublamellosa 39 16 55 1.50% 2.47% 1.70% Composita lewisensis 9 10 19 0.35% 1.54% 0.59%

Table TA11. (Continued)

Composita subquadrata 40 25 65 1.54% 3.85% 2.00% Diaphragmus elegans 16 7 23 0.62% 1.08% 0.71% Eumetria verneuliana 3 0 3 0.12% 0.00% 0.09%

Inarticulata (2) Oehlertella pleurites 2 0 2 0.08% 0.00% 0.06% Orbiculoidea sp. 6 2 8 0.23% 0.31% 0.25%

Stenolaemata (9) Archimedes cf. meekanus 1525 213 1738 58.79% 32.82% 53.59% Eridopora sp. 11 5 16 0.42% 0.77% 0.49% Fenestella sp. 11 1 12 0.42% 0.15% 0.37%

271 Glyptopora punctipora 111 24 135 4.28% 3.70% 4.16%

Lyroporella sp. 1 0 1 0.04% 0.00% 0.03% Polypora sp. 85 14 99 3.28% 2.16% 3.05% Rhombopora sp. 261 171 432 10.06% 26.35% 13.32% Septopora sp. 3 5 8 0.12% 0.77% 0.25% Sulcoretepora sp. 3 1 4 0.12% 0.15% 0.12%

Blastoidea (1) Pentremities elegans 29 12 41 1.12% 1.85% 1.26%

Echinoidea (2) Lepidesthes formosa 1 0 1 0.04% 0.00% 0.03% Archaeocidaris megastylus 0 3 3 0.00% 0.46% 0.09% (isolated spines only)

Crinoidea (16)

Table TA11. (Continued)

Acrocrinus shumardi 1 0 1 0.04% 0.00% 0.03% Agassizocrinus lobatus 1 0 1 0.04% 0.00% 0.03% (juvenile) Anartiocrinus maxvillensis 1 0 1 0.04% 0.00% 0.03% Aphelecrinus mundus 5 0 5 0.19% 0.00% 0.15% Camptocrinus cirrifer 11 1 12 0.42% 0.15% 0.37% Cryphiocrinus girtyi 2 0 2 0.08% 0.00% 0.06% Cymbiocrinus grandis 54 1 55 2.08% 0.15% 1.70% Cymbiocrinus tumidus 11 0 11 0.42% 0.00% 0.34% Dasciocrinus florealis 30 5 35 1.16% 0.77% 1.08% Pentaramicrinus gracilis 25 3 28 0.96% 0.46% 0.86% Phanocrinus bimagnaramus 27 2 29 1.04% 0.31% 0.89% Phanocrinus maniformis 4 0 4 0.15% 0.00% 0.12%

272 Pterotocrinus acutus 10 6 16 0.39% 0.92% 0.49%

Taxocrinus whitfieldi 29 2 31 1.12% 0.31% 0.96% Tholocrinus spinosus 4 0 4 0.15% 0.00% 0.12% Zeacrinites magnoliaeformis 3 0 3 0.12% 0.00% 0.09%

Chondrichthyes (2) Cladodus 1 0 1 0.04% 0.00% 0.03% Venustodus 1 0 1 0.04% 0.00% 0.03%

Grand Total of N 2,594 649 3,243 100.00% 100.00% 100.00%

entire entire 0.05% 1.47% 0.42% 0.21% 0.11% 0.42% 0.42% Entire Entire 14.31% 12.26% Relative Relative Frequency Frequency Landscape

0.00% 0.33% 0.82% 0.49% 0.00% 6.89% 0.49% 0.33% 12.30% Basinal Relative Relative Frequency Frequency

B 0 2 5 3 0 3 2 n 42 75 Basinal Basinal

grained, argillaceous calcarenites argillaceous grained, - 0.00% 4.86% 0.00% 0.00% 0.00% 1.62% 0.00% 4.32% 20.54% Relative Relative Frequency Transitional ), coarse rgillaceous calcilutites and rgillaceous (basinal),

T 0 3 0 8 0 9 0 0 n 38 Transitional Transitional

- ransitional), 0.00% 6.91% 0.53% 0.27% 4.26% 0.00% 3.46% 0.00% 0.00% t Shoal Margin Relative Relative Frequency Frequency -

SM 0 2 1 0 0 0 26 16 13 n Shoal grained calcarenite (shoal calcarenite grained Margin Margin - e calcilutites (

coars Shoal 0.28% 0.00% 0.69% 0.14% 0.55% 0.41% 0.14% 22.96% 18.53% Relative Relative Frequency Frequency

S 2 0 5 1 4 3 1 n 166 134 Shoal

4 2 8 8 1 8 28 (n) 272 233 Number Number -

grained grained grained grained - -

e ), ), Coarse

Fine grained calcarenites interbedded and calcarenites grained -

missouriensis sp.

elative frequency calculations for for the calculations frequency elative Margin), Margin), R -

. and Whole Community and Whole calcilutites (Transitional), (Transitional), calcilutites for the Valley the Valley for Stone Quarry. calcarenite (Shoal calcarenite

calcarenites and interbedded interbedded and calcarenites (Shoal Valley Stone Quarry, Relative Relative Quarry, Stone Valley Frequency for Coars for Frequency grained, argillaceous calcarenites calcarenites argillaceous grained, Argillaceous calcilutites (Basinal), (Basinal), calcilutites Argillaceous Anthracospirifer leidyi Anthracospirifer sublamellosa Cleiothyridina lewisensis Composita Sulcatopinna missouriensis Sulcatopinna (1) Polychaeta? haneyensis Crinicaminus (11) Articulata Scyphozoa? (2) Scyphozoa? Sphenothallus Paraconularia (1) Pelecypoda Demospongea (1) Demospongea fascicularis Belemnospongia (1) Anthozoa coral? tabulate Unidentifiable margin), margin), fine TA12 - landscape Table (shoal

273

Table TA12. (Continued)

Composita subquadrata 410 63 8.71% 215 57.18% 25 13.51% 107 17.54% 21.57% Diaphragmus elegans 26 1 0.14% 3 0.80% 1 0.54% 21 3.44% 1.37% Dielasma sp. 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Eumetria costata 4 2 0.28% 0 0.00% 0 0.00% 2 0.33% 0.21% Eumetria verneuliana 5 2 0.28% 1 0.27% 0 0.00% 2 0.33% 0.26% Orthotetes kaskaskiensis 7 2 0.28% 0 0.00% 0 0.00% 5 0.82% 0.37% Punctospirifer sp. 7 0 0.00% 2 0.53% 0 0.00% 5 0.82% 0.37% Streptorhynchus sp. 1 1 0.14% 0 0.00% 0 0.00% 0 0.00% 0.05%

Inarticulata (2) Lingulipora sp. 2 2 0.28% 0 0.00% 0 0.00% 0 0.00% 0.11% Orbiculoidea sp. 6 1 0.14% 2 0.53% 3 1.62% 0 0.00% 0.32%

Stenolaemata (10)

274 Anisotrypa sp. 3 2 0.28% 0 0.00% 0 0.00% 1 0.16% 0.16%

Archimedes cf. meekanus 17 11 1.52% 6 1.60% 0 0.00% 0 0.00% 0.89% Eridopora sp. 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Fenestella sp. 4 0 0.00% 0 0.00% 1 0.54% 3 0.49% 0.21% Fenestrellina sp. 1 0 0.00% 0 0.00% 1 0.54% 0 0.00% 0.05% Polypora sp. 24 2 0.28% 4 1.06% 5 2.70% 13 2.13% 1.26% Rhombopora sp. 45 13 1.80% 4 1.06% 0 0.00% 28 4.59% 2.37% Septopora sp. 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Thamniscus? 40 16 2.21% 7 1.86% 14 7.57% 3 0.49% 2.10% Unidentifiable encrusting bryozoan 6 5 0.69% 1 0.27% 0 0.00% 0 0.00% 0.32%

Edrioasteroidea (2) Unidentifiable edrioasteroid 6 6 0.83% 0 0.00% 0 0.00% 0 0.00% 0.32% Lepidodiscus laudoni 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05%

Blastoidea (1) Pentremities elegans 89 29 4.01% 8 2.13% 2 1.08% 50 8.20% 4.68%

Table TA12. (Continued)

Echinoidea (1) Archaeocidaris megastylus 54 1 0.14% 0 0.00% 14 7.57% 39 6.39% 2.84%

Ophiuroidea (1) Unidentifiable encrinasterid brittle 34 21 2.90% 2 0.53% 9 4.86% 2 0.33% 2.16% star

Crinoidea (18) Agassizocrinus lobatus 30 3 0.41% 5 1.33% 3 1.62% 19 3.11% 1.58% Anartiocrinus maxvillensis 31 21 2.90% 0 0.00% 3 1.62% 7 1.15% 1.63% Aphelecrinus mundus 25 14 1.94% 7 1.86% 0 0.00% 4 0.66% 1.32% Camptocrinus cirrifer 2 0 0.00% 1 0.27% 1 0.54% 0 0.00% 0.11% Cryphiocrinus girtyi 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Cymbiocrinus grandis 35 15 2.07% 7 1.86% 0 0.00% 13 2.13% 1.84% Cymbiocrinus tumidus 23 8 1.11% 3 0.80% 0 0.00% 12 1.97% 1.21%

275 Dasciocrinus florealis 88 46 6.36% 8 2.13% 12 6.49% 22 3.61% 4.63%

Eupachycrinus boydii 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Linocrinus wachsmuthi 48 36 4.98% 4 1.06% 5 2.70% 3 0.49% 2.52% Pentaramicrinus gracilis 16 2 0.28% 0 0.00% 0 0.00% 14 2.30% 0.84% Phanocrinus bimagnaramus 141 68 9.41% 24 6.38% 26 14.05% 23 3.77% 7.42% Phanocrinus maniformis 1 1 0.14% 0 0.00% 0 0.00% 0 0.00% 0.05% Phacelocrinus longidactylus 1 0 0.00% 1 0.27% 0 0.00% 0 0.00% 0.05% Platycrinites sp. 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Talarocrinus sp. 6 4 0.55% 0 0.00% 2 1.08% 0 0.00% 0.32% Taxocrinus whitfieldi 11 8 1.11% 0 0.00% 0 0.00% 3 0.49% 0.58% Tholocrinus spinosus 1 1 0.14% 0 0.00% 0 0.00% 0 0.00% 0.05%

Chondrichthyes (13) Agassizodus 1 0 0.00% 1 0.27% 0 0.00% 0 0.00% 0.05% Cladodus 3 0 0.00% 1 0.27% 0 0.00% 2 0.33% 0.16% Ctenacanthus 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Deltodopsis 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05%

Table TA12. (Continued)

Deltodus 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Listracanthus? 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Lophodus 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Mesodmodus 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Petrodus 59 0 0.00% 0 0.00% 0 0.00% 59 9.67% 3.10% Psammodus? 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Polyrhizodus 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Taeniodus? 1 0 0.00% 0 0.00% 0 0.00% 1 0.16% 0.05% Venustodus 1 0 0.00% 1 0.27% 0 0.00% 0 0.00% 0.05%

Grand Total: 1,894 723 100.00% 376 100.00% 185 100.00% 610 100.00% 100.00%

276

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293 VITA Author’s Name: Ann Well Harris

Place of Birth: Manchester, Kentucky

Education:  M.S. Geosciences, 2009, Eastern Kentucky University  B.S. Geology, 2007, Eastern Kentucky University

Work Experience:  Adjunct Instructor, 2012 − present, University of Pikeville  Adjunct Instructor, 2011 − present, Hazard Community and Technical College  Adjunct Instructor, 2010 – present, Eastern Kentucky University  GIS Assistant, 2006 − present, Cumberland Valley Area Development District

Awards:  The Manchester Enterprise, Best of the Best Annual Award, Best EKU Professor Award, 2014, 2015, 2016, and 2017.  Graduate Research Competition Award, Kentucky Academy of Science Annual Meeting (KAS), 1st place in geology (November, 2015), 3rd pace in geology (October, 2012), 2nd place in geology (November, 2011), 1st place in geology (November, 2010).

Abstracts/Publications:

 Harris, A.W., and Lierman, R.T., 2011, Examination of a Tabulate Coral Biostrome in the Bull Fork Formation (Ordovician), Bath County, Kentucky: Geological Society of America Abstracts with Programs, v. 43, No. 2, p. 20.  Harris, A.W., and Lierman, R.T., 2011, Examination of a Tabulate Coral Biostrome in the Bull Fork Formation (Ordovician), Bath County, Kentucky: The Professional Geologist, v. 48, No. 4, p. 46–51.  Harris, A.W., and Ettensohn, F.R., 2015, A History and Census Report from two Late Mississippian bryozoan/echinoderm faunules, Powell County, East-Central Kentucky: Geological Society of America Abstracts with Programs, v. 47, No. 2, p.81.  Harris, A.W., and Ettensohn, F.R., 2015, A Comparison of Environmental Constraints on Late Mississippian (Chesterian) Echinoderm faunules from East- Central Kentucky, U.S.A: Geological Society of America Abstracts with Programs, v. 47, No. 7, p.765.  Harris, A.W., and Ettensohn, F.R., 2017, Population Analysis of a Late Mississippian (Chesterian) Echinoderm Faunule Across Four Contiguous Depositional Environments, Carter County, East-Central Kentucky: Geological Society of America Abstracts with Programs, v. 49, No. 2.

294  Harris, A.W., Ettensohn, F.R., and Carnahan-Jarvis, J., 2018, A statistical approach to understanding diversity in an Upper Mississippian (Chesterian) echinoderm-rich unit across four contiguous lithofacies, Carter County, northeastern Kentucky: Geological Society of America Abstracts with Programs, v. 50, No. 3, ISSN 0016-7592.

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