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By Christopher Boyd a Thesis Submitted to the School of Graduate THREE-DIMENSIONAL TOMOGRAPHIC RECONSTRUCTION OF AN EXCEPTIONALLY WELL PRESERVED ICHNOLOGICAL ASSEMBLAGE FROM THE STAINMORE FORMATION, CARBONIFEROUS, UK. By Christopher Boyd A thesis submitted to the School of Graduate Studies in partial fulfilment of the requirements for the degree of Doctor of Philosophy Department of Earth Science Memorial University of Newfoundland May, 2016 St. John’s Newfoundland and Labrador THREE DIMENSIONAL TOMOGRAPHIC RECONSTRUCTION OF AN EXCEPTIONALLY WELL PRESERVED ICHNOLOGICAL ASSEMBLAGE ABSTRACT This Ph.D. thesis addresses current issues with ichnotaxonomic practice, and characterizes an exceptionally well preserved ichnological assemblage from the Carboniferous Stainmore Formation, Northumberland, United Kingdom. Samples were collected from closely localized float representative of various units throughout the succession, which was deposited in a storm-dominated marine shoreface. Three dominant ichnotaxa were selected for three-dimensional morphological analysis due to their complicated morphology and/or unclear taxonomic status: 1) Dactyloidites jordii isp. nov.; 2) Beaconites capronus, and; 3) Neoeione moniliformis comb. nov. Using serial grinding and photography, these ichnotaxa were ground and modelled in true colour. High-resolution models of three taxa produced in this study are the basis of the first complete three-dimensional consideration of the traces, and forms the basis for refined palaeobiological and ethological analysis of these taxa. Dactyloidites jordii isp. nov. is a stellate to palmate burrow composed of numerous long, narrow rays that exhibit three orders of branching arranged into tiered galleries radiating from a central shaft. It is considered to be the feeding structure produced by a vermiform organism. Beaconites capronus is a winding trace with distinctly chevron-shaped, meniscate backfill demonstrated herein to backfill the vertical shafts associated with its burrows in a comparable fashion to the horizontal portion of the burrow. This lack of a surface connection would result in the trace making organism being exposed to low-oxygen porewater. Coping with this porewater dysoxia could be approached by burrowing organisms in a number of ways: 1) revisiting the sediment-water interface; 2) creating periodic shafts; or 3) employing anaerobic metabolism. Neoeione moniliformis was originally introduced as Eione moniliformis, however, the genus Eione Tate, 1859 is a junior homonym of Eione Rafinesque, 1814. This led to the transfer of Eione moniliformis to Parataenidium. Through careful examination and three-dimensional characterization of topotypes, the transfer to Parataenidium moniliformis is demonstrated herein to be problematic, as Parataenidium refers to primarily horizontal burrows with two distinct layers and Eione moniliformis is composed of one distinct level. As such, the new ichnogenus Neoeione is created to accommodate Neoeione moniliformis. ii THREE DIMENSIONAL TOMOGRAPHIC RECONSTRUCTION OF AN EXCEPTIONALLY WELL PRESERVED ICHNOLOGICAL ASSEMBLAGE ACKNOWLEDGEMENTS “Ichnoledgements” Firstly, I would like to thank my first teacher, Ms Betty Bissell: my kindergarten teacher. Thank you for the continued support and influence you have had on my life and for encouraging me to play in the sandbox. I would also like to express my gratitude to my supervisor, Dr. Duncan McIlroy. I would like to thank you for the opportunity, guidance, and most of all for allowing me the freedom to decide which direction to take the project. Your support and overall passion for science have made me a better scientist. A special thank you goes to Dr. Suzanne Dufour and Dr. Richard Callow, my supervisory committee. I have appreciated your ideas and guidance throughout this project. I would like also to thank Edgars Rudzitis, Dr. Liam Herringshaw, Dr. Małgorzata Bednarz, and Dr. Michael Garton for your technical assistance through the course of my research. Whether you have helped me with discussions of taxonomy, sample processing, technical glitches in the modelling software, or wielding the gas powered saw to trim samples, your help and your smiles throughout have been appreciated. To Jill Davis of the British Museum of Natural History, Nathalie Djan-Chékar of the Rooms Provincial Museum, and Dr. Steven Holland of the University of Georgia, your assistance with photographing, processing, and searching for the requisite type specimens has been essential for the progress of my study. To my family, I can’t thank you enough for all the years of support and encouragement you have provided me. I know you understand how busy this time has been, but I hope you also know how much I have enjoyed every moment with you all, and how much I wish I could have attended more Sunday dinners. Mom and Dad, thank you for your advice, encouragement, and love (Oh and dinners to go!). To my little guy Zach, thank you for all the play time and dinosaur-talk, and for being a source of some essential respite. To my best friend, Robyn. You have made the hardest, most challenging days a pleasure simply by being by my side. Your compassion and tenderness is only matched by your incredible intellect. It is amazing to have you with me, even just to sip wine and talk science with. You’ve been the inspiration for some of my best ideas, in particular asking to marry you! iii THREE DIMENSIONAL TOMOGRAPHIC RECONSTRUCTION OF AN EXCEPTIONALLY WELL PRESERVED ICHNOLOGICAL ASSEMBLAGE TABLE OF CONTENTS Abstract ii Acknowledgements iii Table of Contents iv List of Tables vii List of Figures vii List of Appendices ix Co-Authorship Statement x CHAPTER ONE – ICHNOLOGY AND ICHNOTAXONOMY: INTRODUCTION AND 1-1 OVERVIEW 1. Introduction to ichnology and literature review 1-1 1.1 History of ichnology 1-2 1.2 Introduction to ichnotaxonomy 1-5 1.3 History of ichnotaxonomy 1-6 1.4 International Code of Zoological Nomenclature 1-8 1.4.1 Scope and exclusions 1-8 1.4.2 Concerns 1-9 1.5 Ichnotaxa as compared with biotaxa 1-13 1.6 Description/characterization methods 1-17 1.6.1 Morphometrics 1-19 1.6.2 Valid Ichnotaxobases 1-19 1.6.3 Ichnotaxobase hierarchy 1-21 1.7 Near burrow sedimentary structures 1-24 1.7.1 Structures that reflect ethology 1-25 1.7.2 Non-ethological sedimentary features 1-31 2. Thesis Objectives 1-32 2.1 Aims and objectives of CHAPTER TWO 1-33 2.2 Aims and objectives of CHAPTER THREE 1-34 2.3 Aims and objectives of CHAPTER FOUR 1-34 iv THREE DIMENSIONAL TOMOGRAPHIC RECONSTRUCTION OF AN EXCEPTIONALLY WELL PRESERVED ICHNOLOGICAL ASSEMBLAGE 2.4 Aims and objectives of CHAPTER FIVE 1-35 3. Methodology 1-35 3.1 Serial grinding and photographic digital modelling 1-36 3.1.1 Improvements on the base techniques 1-38 3.2 Field assessment and sample collection/characterization 1-39 4. Study Area 1-39 5. References 1-41 CHAPTER TWO – THREE-DIMENSIONAL MORPHOLOGY AND PALAEOBIOLOGY 2-1 OF DACTYLOIDITES JORDII ISP. NOV Abstract 2-2 1. Introduction 2-3 2. Methodology 2-7 3. Systematic ichnology 2-9 4. Palaeobiology 2-18 5. Conclusions 2-22 6. Acknowledgements 2-24 7. References 2-24 CHAPTER THREE – THREE-DIMENSIONAL MORPHOLOGY OF BEACONITES 3-1 CAPRONUS FROM NORTHEAST ENGLAND Abstract 3-2 1. Introduction 3-3 2. Geologic Setting 3-5 3. Methodology and Modelling 3-6 4. Results 3-8 4.1 Burrow morphology 3-8 5. Systematic ichnology 3-14 6 Discussion 3-17 6.1 Palaeobiology 3-17 6.2 Nutrition 3-18 v THREE DIMENSIONAL TOMOGRAPHIC RECONSTRUCTION OF AN EXCEPTIONALLY WELL PRESERVED ICHNOLOGICAL ASSEMBLAGE 6.3 Locomotion 3-20 6.4 Vertical shafts 3-21 7. Conclusion 3-22 9. Acknowledgements 3-23 10. References 3-23 CHAPTER FOUR – THE STRUCTURE, MORPHOLOGY, AND METHOD OF 4-1 FORMATION OF NEOEIONE MONILIFORMIS IGEN. NOV., COMB. NOV. Abstract 4-2 1. Introduction 4-3 2. Methodology and Modelling 4-7 3. Taxonomic background: from Eione to Parataenidium 4-8 4. Descriptive Ichnology 4-10 5. Systematic Ichnology 4-17 6. The terminal portion of a probable N. moniliformis 4-19 7. Palaeobiology 4-22 8. Evidence from a hypidiomorphic Neoeione moniliformis 4-23 9. Conclusion 4-25 10. Acknowledgements 4-26 11. References 4-27 CHAPTER FIVE – SYSTEMATIC ICHNOLOGY OF THE STAINMORE FORMATION 5-1 NEAR HOWICK, NORTHUMBERLAND, UNITED KINGDOM Abstract 5-2 1. Introduction 5-3 1.1 Geological and palaeoenvironmental setting 5-3 2 Systematic Ichnology 5-6 3 Conclusions 5-34 4. Acknowledgements 5-36 5. References 5-36 vi THREE DIMENSIONAL TOMOGRAPHIC RECONSTRUCTION OF AN EXCEPTIONALLY WELL PRESERVED ICHNOLOGICAL ASSEMBLAGE CHAPTER SIX – PROJECT SUMMARY AND CONCLUSIONS 6-1 1. Summary 6-1 1.1 Review of objectives 6-2 1.2 Review of the methodology 6-3 1.3 Outcomes of Chapter Two 6-4 1.4 Outcomes of Chapter Three 6-6 1.5 Outcomes of Chapter Four 6-7 1.6 Outcomes of Chapter Five 6-8 2. Concluding Statement 6-8 3. References 6-9 LIST OF TABLES CHAPTER ONE 1-1 Table Major contemporary species concepts and their key differentiating 1-16 1-1: properties. Table Various works of organisms which are either considered valid or 1-18 1-2: invalid traces. Table Methods of characterizing Ichnofamilies. 1-24 1-3: Table Criteria for distinguishing ethological versus non-ethological 1-32 1-4: structures. CHAPTER TWO 2-1 Table Dactyloidites jordii morphometrics 2-14 2-1: LIST OF FIGURES CHAPTER ONE 1-1 Figure 1-1: Illustration of a hypothetical compound trace. 1-13 Figure 1-2: Graded bedding in Tasselia ordamensis. 1-29 Figure 1-3: Skolithos burrow compared with Monocraterion. 1-29 Figure 1-4: Collapse cones in relation to Ophiomorpha and Thalassinoides. 1-30 Figure 1-5: Modern collapse cones created by Neotrypaea californiensis. 1-30 vii THREE DIMENSIONAL TOMOGRAPHIC RECONSTRUCTION OF AN EXCEPTIONALLY WELL PRESERVED ICHNOLOGICAL ASSEMBLAGE CHAPTER TWO 2-1 Figure 2-1: Map of field location and sample location 2-5 Figure 2-2: Sedimentary field log of the Stainmore Formation located near the 2-6 town of Howick Figure 2-3: Digital model of Dactyloidites jordii isp.
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