Durham E-Theses The palaeobiology of the panderodontacea and selected other euconodonts Sansom, Ivan James How to cite: Sansom, Ivan James (1992) The palaeobiology of the panderodontacea and selected other euconodonts, Durham theses, Durham University. Available at Durham E-Theses Online: http://etheses.dur.ac.uk/5743/ Use policy The full-text may be used and/or reproduced, and given to third parties in any format or medium, without prior permission or charge, for personal research or study, educational, or not-for-prot purposes provided that: • a full bibliographic reference is made to the original source • a link is made to the metadata record in Durham E-Theses • the full-text is not changed in any way The full-text must not be sold in any format or medium without the formal permission of the copyright holders. Please consult the full Durham E-Theses policy for further details. Academic Support Oce, Durham University, University Oce, Old Elvet, Durham DH1 3HP e-mail: [email protected] Tel: +44 0191 334 6107 http://etheses.dur.ac.uk The copyright of this thesis rests with the author. No quotation from it should be pubHshed without his prior written consent and information derived from it should be acknowledged. THE PALAEOBIOLOGY OF THE PANDERODONTACEA AND SELECTED OTHER EUCONODONTS Ivan James Sansom, B.Sc. (Graduate Society) A thesis presented for the degree of Doctor of Philosophy in the University of Durham Department of Geological Sciences, July 1992 University of Durham. 2 DEC 1992 Contents CONTENTS CONTENTS p. i ACKNOWLEDGMENTS p. viii DECLARATION AND COPYRIGHT p. ix ABSTRACT p. xii Chapter 1 INTRODUCTION M A brief history of discovery and the development of conodont taxonomy p. 1 The Granton conodont animals p. 2 The Waukesha Panderodus animal p. 3 Conodont phylogeny p. 4 Conodont palaeobiology p. 5 Chapter 2 THE APPARATUS ARCHITECTURE OF PANDERODUS p. 6 2.1 INTRODUCTION p. 6 2.2 GENERIC AND SPECIES CONCEPTS IN PANDERODUS p. 8 23 THE COMPOSITION OF PANDERODUS APPARATUSES p. 13 2.4 CLUSTER DESCRIPTIONS p. 15 2.4.1 Waukesha Bedding Plane Assemblage p. 16 2.4.2 Nekezseny Cluster p. 18 Contents 2.43 Podolia Cluster p. 19 2.4.4 Shandong Cluster p. 21 2.4.5 Isolated Waukesha Clusters p. 21 2.5 CLUSTER CORRELATION p. 24 2.6 HOMOLOGY BETWEEN CONIFORM GENERA p. 26 2.6.1 The integrity of the genus Panderodus p. 26 2.6.2 Applicability of the Panderodus apparatus model to other Panderodontid lineages p. 27 Belodina compressa (Branson and Mehl) p. 28 Parapanderodus Stouge p. 31 2.6.3 Applicability of the Panderodus apparatus model to Non- Panderodontidae coniforms p. 34 Coelocerodontus Ethington, 1959 p. 34 Besselodus Aldridge, and Dapsilodus Cooper p. 36 2.7 IMPLICATIONS FOR PANDERODONTID SUPERGENERIC CLASSIFICATION p. 39 Chapter 3 A PRELIMINARY ATTEMPT TO UNIFY CONODONT LOCATIONAL NOMENCLATURE pJ3 3.1 INTRODUCTION p. 43 3.2 CONIFORM APPARATUSES p. 43 3.3 RAMIFORM-PECTINIFORM APPARATUSES p. 45 33.1 Order Ozarkodinida Dzik 1976 p. 45 33.2 Order Prioniodontida Dzik 1976 p. 47 3.4 COMPARISON BETWEEN OZARKODIND) AT^ PANDERODONTID APPARATUSES p. 49 Contents 3.5 EVOLUTIONARY DEVELOPMENT OF APPARATUS ARCHITECTURE p. 52 Chapter 4 A FUNCTIONAL INTERPRETATION OF THE PANDERODUS APPARATUS p^ 4.1 INTRODUCTION p. 56 4.2 HISTORICAL REVIEW p. 57 43 A FUNCTIONAL MODEL FOR THE PANDERODUS APPARATUS p. 65 4.4 POSSIBLE FUNCTIONAL ANALOGUES p. 67 4.4.1 Comparison with the chaetognath grasping array p. 67 4.4.2 Comparison with the lingual apparatus of myxinoids p. 68 4.4.3 Comparison with the oral apparatuse of the petromyzontids p. 71 4.5 THE FEEDING APPARATUSES OF EARLY 'AGNATHANS' p. 73 4.6 CONCLUSIONS p. 75 Chapter 5 THE HISTOLOGY OF SELECTED CONODONT ELEMENTS EJZ 5.1 INTRODUCTION p. 77 5.1.1 Protoconodonts, Paraconodonts and Euconodonts p. 77 5.2 VERTEBRATE ODONTOGENIC TISSUES p. 82 Enamel and Enameloid p. 82 Dentine p. 83 Bone p. 83 HI Contents Aspidin p. 83 Cartilage p. 84 53 THE DEVELOPMENTAL SEQUENCE OF VERTEBRATE ODONTOGENIC TISSUES p. 84 5.4 THE HARDING SANDSTONE VERTEBRATES p. 86 5.5 HISTORICAL REVIEW OF CONODONT HISTOLOGICAL WORK p. 92 5.5.1 Technological Developments in the Study of Conodont Mineralised Tissues p. 92 5.5.2 Previous Interpretations of Conodont Histology p. 93 5.6 METHODOLOGY OF CONODONT HISTOLOGICAL STUDIES p.lOO 5.7 HISTOLOGICAL DESCRIPTION AND INTERPRETATION OF CONODONT HARD TISSUES p. 102 5.7.1 Lamellar Crown Tissue p.l03 5.7.2 White Matter p.l04 5.7 J Basal Body p. 105 5.8 THE DISTRIBUTION OF VERTEBRATE TISSUES IN CONODONT ELEMENTS p.l06 5.8.1 The Family Panderodontidae p.l06 Panderodus (Ethington) p.l06 Parapanderodus striatus (Graves and Ellison) p.l09 5.8.2 Non-Panderodontids p.llO Order Proconodontida Sweet, 1988 p.llO Cordylodus Pander p. 110 Eoconodontus Miller p. 111 Order Ozarkodinida Dzik, 1976 p.ll2 Ozar^oJ/na (Branson and Mehl) p.ll2 Order Belodellida Sweet, 1988 p.l 12 IV Contents Wapsilodus Cooper p. 113 Order Unknown p. 113 Pseudooneotodus Drygant p. 113 5.9 DISCUSSION p.ll4 Chapter 6 CONODONT RELATIONSHIPS AND A REINTERPRETATION OF CHORDATE PHYLOGENY p^ 6.1 INTRODUCTION p.l20 6.2 CLADISTIC TERMINOLOGY p. 120 63 TAXONOMIC UNITS p.l22 Conodonts (Late Cambrian - end Triassic) p.l22 Late Cambrian - Early Ordovician Problematica p.l23 Urochordates p.l25 Cephalochordates p. 127 Myxinoids p.l28 Astraspis (Caradoc, Late Ordovician) p.l29 Eriptychius (Caradoc, Late Ordovician) p.l3l 3rd unnamed vertebrate from the Harding Sandstone (Caradoc, Late Ordovician) p.l32 Other Ordovician 'agnathans' p.l32 Heterostracans (Early Silurian - Late Devonian) p.l34 Petromyzontids p.l35 Osteostracans (Early Silurian - Late Devonian) p.l36 Gnathostomes (Early Silurian - Present) p. 137 Post-Ordovician problematic vertebrates p.l38 6.4 CHARACTERS p. 139 Contents 6.4.1 Histological Characters p.l40 Cartilaginous endoskeleton p.l41 Dermal ossification p.l42 Superficial cap p. 142 Dentine p.l43 Cellular - acellular bone p.l43 Aspidin p.l44 Honeycombed middle layer p.l44 Perichondral Bone and Endochondral Bone p.l45 6.4.2 Features relating to the bodyplan p. 145 Somites p.l45 Heterocercal Tail p.l45 Paired fins p.l46 Bite p. 146 One paired Branchial opening p. 147 Slanted Gill Openings p.l47 Nerves IX and X issuing from braincase p.l48 6.5 RESULTS AND DISCUSSION p.l48 Chapter 7 SYSTEMATIC PALAEONTOLOGY p.l61 7.1 GENERAL CONSIDERATIONS p.l61 7.2 TERMINOLOGY p. 162 Genus Panderodus Ethington, 1959 p.l65 Panderodus bergstromi Sweet, 1979 p.l70 Panderodus breviusculus Barnes, 1977 p. 171 Panderodus equicostatus (Rhodes, 1953) p. 172 VI Contents Panderodus feulneri (Glenister, 1957) p.l77 Panderodus gracilis (Branson and Mehl, 1933) p. 181 Panderodus langkawiensis (Igo and Koike, 1967) p. 185 Panderodus levigatus (Nowlan and McCracken, 1988) p. 188 Panderodus panderi (Stauffer, 1940) p. 189 Panderodus rhamphoides Nowlan and McCracken (in Nowlan et a/.), 1988 p. 194 Panderodus staufferi (Branson, Mehl and Branson, 1951) p.l95 Panderodus sulcatus (FShrseus, 1966) p. 199 Panderodus unicostatus (Branson and Mehl, 1933) p.201 Chapter 8 CONCLUSIONS AND FUTURE RESEARCH p^ References p.209 Plates p.237 Appendix 1: SAMPLE LOCATION AND PREPARATION p.257 Appendix 2: RESIN SUPPLIERS p.262 Vll Acknowledgements Drs Howard A Armstrong (Durham) and M. Paul Smith (Birmingham) were responsible for the initiation and supervision of this project. Fd like to thank H.AA for introducing me to the bizarre world that is conodontology, and both H.AA and M.P.S. for trying to constrain my wildest (scientific) fantasies. I also acknowledge Prof. Joe Cann and Dr. Mick Jones for their permission to use facilities within the Department of Geology at Newcastle upon Tyne, and Prof. Roger Searle (Durham) for allowing me to freeload in the Department of Geological Sciences at Durham. During the course of this thesis, I have benefited from encouragement, advice and unpublished information from the following: Dr. Richard J. Aldridge (Leicester), for allowing me to walk off with his most pri?ed Panderodus specimens; Dr. Lennart Jeppsson (Lund), for actually believing Panderodus has its uses; Dr. Moya M. Smith (Guy's Hospital), who occasionally baffled me with vertebrate embryology; Dr. Graham A Young (Manitoba), for letting me dissolve some very pretty corals from Gotland; and the large number of Panderers with whom I have been in regular contact, notably Dr. Rod Norby (Illinois State Geological Survey), who supplied the Waukesha clusters, and Anita Lofgren (Lund), who gave me the P. sulcatus samples. Td also like to say 'Cheers' to all those who attended Lacon I in Bolivia and Argentina, and made that trip so much fun. Technically, this thesis would not have been possible without the help of a large number of people at both Newcastle and Durham. I would like to thank Trevor Whitfield, Brian Tuffs, Brian Richardson and Len Rhodes for their help whilst I was resident in Newcastle; and Dave Asbery, George Randall, Alan Carr, Gerry Dresser and Karen Atkinson for giving me enough rope in Durham. Angus Parker and Trevor Booth (Newcastle Medical School) are singled out for their excellent S.E.M. service, and for pointing me in the right directions during the histological work. Thanks must also go to friends and colleagues from Newcastle and Durham, notably Doug Forbes, Cherry Walker, Ian Billing and Tim Alsop, who have all dragged me kicking and screaming to a suitable watering hole when it seemed Fd die of dehydration. Finally, and by no means least, Td like to express a debt of extreme gratitude to Jane Keeton for making the butties, proof-reading the thesis, and putting up with me whilst I've been a pain in the butt.