MARINE VERTEBRATE COMMUNITIES FROM THE CISURALIAN EPOCH (PERMIAN PERIOD) OF CENTRAL NORTH AMERICA A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy By RYAN C SHELL B.S., University of Arkansas, 2013 2020 Wright State University COPYRIGHT BY RYAN C SHELL 2020 WRIGHT STATE UNIVERSITY GRADUATE SCHOOL September 11, 2020 I HEREBY RECOMMEND THAT THE DISSERTATION PREPARED UNDER MY SUPERVISION BY Ryan C Shell ENTITLED Marine Vertebrate Communities From The Cisuralian Epoch (Permian Period) Of Central North America BE ACCEPTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF Doctor of Philosophy _______________________ _______ Charles Ciampaglio, Ph.D. Dissertation Director _______________________________ Donald Cipollini, Ph.D Program Director _______________________________ Barry Milligan, Ph.D. Interim Dean of the Graduate School Committee on Final Examination: _____________________________ Charles Ciampaglio, Ph.D ____________________________ Rebecca Teed, Ph.D ____________________________ Stephen Jacquemin, Ph.D. ____________________________ David Schmidt, Ph.D ____________________________ Alexander Ivanov, Ph.D ABSTRACT Shell, Ryan C. Ph.D. Environmental Sciences Ph.D. Program, Wright State University, 2020. Marine vertebrate communities from the Cisuralian Epoch (early Permian Period) of central North America. Marine vertebrates from the Cisuralian Epoch early Permian Period) are rare in the global fossil record. In particular, species-rich faunas from individual localities are poorly represented, with single occurrence localities that do not fully capture faunal tends in marine vertebrate ecology across this interval. This dissertation helps to close this gap by reporting marine vertebrate fossils from five localities across central North America, containing four to 15 unequivocal vertebrate taxa, as well as a new single occurrence locality near the Carboniferous-Permian boundary. This survey includes representatives from the first three of the four Cisuralian faunal stages, and is the result of new field and lab investigations of vertebrate macro and microremains. An analysis of the number and identity of these fossils at the ordinal level indicates marine vertebrate communities during this interval were likely dominated by hybodontiform and ctenacanthiform chondrichthyans. Other groups such as the Petalodontiformes, Symmoriiformes, Eugeneodontiformes, and Osteichthyes were major components to these many of these assemblages as well. Orodontiforms, neoselachians, bransonelliforms, and cochliodontiforms also occurred in smaller numbers at various points during the Cisuralian Epoch. iv TABLE OF CONTENTS 1. INTRODUCTION……………………………………………………………1 1.1 CISURALIAN MARINE VERTEBRATE ECOLOGY AND THE ASSOCIATED RESEARCH…………………………………………...…1 1.2 RESEARCH GAPS………………...…………………....……….…..........3 1.3 INTRODUCTION OF STUDY AREAS…………………………………..7 1.4 HYPOTHESIS…………………………………………………………..…12 1.5 GENERAL PROCEDURES………………………………………………14 1.6 NOTE ON STRUCTURE OF THIS DISSERTATION…………………..16 1.7 LITERATURE CITED………………………………………………...….16 2. SHARK REMAINS NEAR THE CARBONIFEROUS-PERMIAN BOUNDARY IN THE HARPERSVILLE FORMATION OF TEXAS……………………...22 2.1 INTRODUCTION……………………………………………………..….22 2.2 METHODS…………………………………………………...……..…….24 2.3 RESULTS…………………………………………………………………24 2.3.1 SYSTEMATIC PALEONTOLOGY………………………………..26 2.4 DISCUSSION……………………………………………………………..28 2.5 LITERATURE CITED…………………………………………………....29 3. A VERTEBRATE FAUNA FROM THE NEVA FORMATION (ASSELIAN AGE) OF EASTERN KANSAS…………………………………………………….31 3.1 INTRODUCTION……………………………………………………...….31 v 3.2 METHODS………………………………………………………………..39 3.3 GEOLOGIC SETTING………………………………………………..…..39 3.4 RESULTS……………………………………………………………..…...43 3.4.1 SYSTEMATIC PALEONTOLOGY………………………………....43 3.5 CONCLUSIONS………………………………………………………........62 3.6 DISCUSSION……………………………………………………………….64 3.7 LITERATURE CITED…………………………………………………...…65 4. ADDITIONAL VERTEBRATES FROM THE WREFORD LIMESTONE (SAKMARIAN STAGE) OF SOUTHERN KANSAS………………………71 4.1 INTRODUCTION…………………………………………………...……..71 4.2 GEOLOGIC SETTING…………….………………………………...…….74 4.3 METHODS……………………………………………………………..….77 4.4 RESULTS……………………………………………………………..……78 4.4.1 SYSTEMATIC PALEONTOLOGY………….……………………78 4.5 CONCLUSIONS……………………………………………………………93 4.6 LIERATURE CITED………………………………………………….……94 5. A BREIF HISTORY OF STUDY AND NOTE ON THE LUEDERS FORMATION………………………………………………………………….99 5.1 HISTORY OF STUDY IN THE LUEDERS FORMATION…………..…..99 5.2 A NOTE ON CHAPTERS 6, 7 AND 8…………………………………...101 5.3 LITERATURE CITED………………………………………………..…..104 6. A FOSSIL MOLLUSCAN FAUNA FROM THE LOWER LUEDERS FORMATION OF NORTH-CENTRAL TEXAS………………………....107 vi 6.1 ABSTRACT …………………………………………………………..…107 6.2 INTRODUCTION………………………………………………………..107 6.3 GEOLOGIC SETTING AND STRATIGRAPHY…………………….…109 6.4 METHODS……………………………………………………………….115 6.5 RESULTS………………………………………………………………...118 6.5.1 SYSTEMATIC PALEONTOLOGY…………………………..…119 6.5.2 FAUNAL LIST…………………………………………………..133 6.5.3 BIOSTRATIGRAPHY…………………………………………..134 6.6 DISCUSSION……………………………………………………….......136 6.7 ACKNOWLEDGEMENTS……………………………………………..137 6.8 REFERENCES CITED………………………………………………….137 7. A MARINE VERTEBRATE FAUNA FROM THE EARLY PERMIAN (ARTINSKIAN) LUEDERS FORMATION OF NORTH-CENTRAL TEXAS, U.S.A………………………………………………………………………...146 7.1 KEYWORDS……………………………………………………………146 7.2 ABSTRACT……………………………………………………………..146 7.3 INTRODUCTION……………………………………………….………146 7.4 GEOLOGIC SETTING..………………………………………….……..149 7.5 METHODS……………………………………………………….…...…153 7.6 INSTITUTIONAL ABBREVIATIONS…………………………..……..154 7.7 SYSTEMATIC PALEONTOLOGY………………………………...…...155 7.8 FAUNAL LIST...........................................................................................172 7.9 DISCUSSION.............................................................................................175 vii 7.10 LITERATURE CITED…………………………………………………179 8. AN OFF-SHORE MARINE VERTEBRATE FAUNA FROM THE EARLY PERMIAN (ARTINSKIAN) LUEDERS FORMATION OF NORTH-CENTRAL TEXAS, U.S.A………………………………………………………………185 8.1 KEYWORDS…………………………………………………………….185 8.2 ABSTRACT……………………………………………………………...185 8.3 INTRODUCTION………………………………………………………..186 8.4 METHODS……………………………………………………………….186 8.5 GEOLOGIC SETTING…………………………………………………..187 8.6 RESULTS………………………………………………………………...192 8.6.1 INSTITUTIONAL ABBREVIATIONS…………………………192 8.6.2 SYSTEMATIC PALEONTOLOGY…………………………….192 8.7 DISCUSSION……………………………………………………………201 8.8 LITERATURE CITED…………………………………………………..203 9. CONCLUSIONS AND SUGGESTIONS FOR FUTURE RESEARCH..206 9.1 NEWLY REPORTED, HIGH SPECIES RICHNESS, CISURALIAN MARINE VERTEBRATE LOCALITIES IN SUMMARY………………………..206 9.2 COMPLEX MARINE VERTEBRATE SITES FROM THE CISURALIAN OF NORTH AMERICA AND THEIR TEMPORAL CONTEXT………….208 9.3 TOWARDS A MORE COMPREHENSIVE DESCRIPTION OF COMMUNITY COMPOSITION DURING THIS TIME………………………………..221 9.4 FUTURE RESEARCH………………………………………………….223 viii LIST OF FIGURES Figure 1.1…………………………………………………………………………………..2 Figure 1.2………………………………………………………………………………….5 Figure 1.3…………………………………………………………………………………..7 Figure 1.4…………………………………………………………………………………10 Figure 1.5…………………………………………………………………………………11 Figure 2.1…………………………………………………………………………………26 Figure 2.2…………………………………………………………………………………28 Figure 3.1…………………………………………………………………………………40 Figure 3.2…………………………………………………………………………………41 Figure 3.3…………………………………………………………………………………43 Figure 3.4…………………………………………………………………………………44 Figure 3.5…………………………………………………………………………………44 Figure 3.6…………………………………………………………………………………46 Figure 3.7…………………………………………………………………………………47 Figure 3.8…………………………………………………………………………………52 Figure 3.9…………………………………………………………………………………54 Figure 3.10..………………………………………………………………………………55 ix Figure 3.11..………………………………………………………………………………57 Figure 3.12..………………………………………………………………………………58 Figure 3.13..………………………………………………………………………………59 Figure 3.14..………………………………………………………………………………61 Figure 3.15………………………………………..………………………………………62 Figure 4.1…………………………………………………………………………………76 Figure 4.2…………………………………………………………………………………77 Figure 4.3…………………………………………………………………………………80 Figure 4.4…………………………………………………………………………………82 Figure 4.5……………………………………………………………………………...…..84 Figure 4.6……………………………………………………………………………...…..85 Figure 4.7………………………………………………………………………………….86 Figure 4.8………………………………………………………………………………….88 Figure 4.9…………………………………………………………………………….……89 Figure 4.10..……………………………………………………………………………….91 Figure 4.11..………………………………………………………………………………92 Figure 5.1……………………………………………………………………………….101 Figure 5.2……………………………………………………………………………….103 x Figure 6.1………………………………………………………………………………111 Figure 6.2………………………………………………………………………………113 Figure 6.3………………………………………………………………………………116 Figure 6.4....……………………………………………………………………………119 Figure 6.5………………………………………………………………………………121 Figure 6.6………………………………………………………………………………123 Figure 6.7………………………………………………………………………………125 Figure 6.8………………………………………………………………………………127 Figure 6.9………………………………………………………………………………128 Figure 6.10..……………………………………………………………………………131 Figure 6.11..……………………………………………………………………………132 Figure 6.12..……………………………………………………………………………135 Figure 7.1………………………………………………………………………………148 Figure 7.2………………………………………………………………………………150 Figure 7.3………………………………………………………………………………151 Figure 7.4………………………………………………………………………………152 Figure 7.5………………………………………………………………………………157 Figure 7.6………………………………………………………………………………158 xi Figure 7.7………………………………………………………………………………161 Figure 7.8………………………………………………………………………………162 Figure 7.9………………………………………………………………………………164 Figure 7.10..……………………………………………………………………………165 Figure 7.11..……………………………………………………………………………166 Figure 7.12..……………………………………………………………………………167 Figure