A MULTI-METHOD STUDY ON TRIASSIC CONODONTS Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften an der Karl-Franzens-Universität Graz Institut für Erdwissenschaften Vorgelegt von Yanlong Chen Graz, Oktober 2015 CONTENTS PREFACE 1 ACKNOWLEDGMENTS 2 ABSTRACT 3 CHAPTER 1 6 INTRODUCTION TO THE END-PERMIAN MASS EXTINCTION AND THE FOLLOWING RECOVERY, THE TRIASSIC PALEOGEOGRAPHY AND THE STATUS OF TRIASSIC CONODONT STUDY 1.1 The end-Permian mass extinction and the following recovery 6 1.2. The paleogeography of the study areas 9 1.3. Introduction to Triassic conodonts 10 1.3.1. Shape categories of conodonts 11 1.3.2. Conodont biostratigraphy of Triassic 13 1.4. References 14 CHAPTER 2 21 SIZE VARIATION OF CONODONTS DURING THE SMITHIAN–SPATHIAN (EARLY TRIASSIC) GLOBAL WARMING EVENT Abstract 21 2.1. Introduction 22 2.2. Geological setting 22 2.3. Method and materials 23 2.4. Results 25 2.5. Discussion 27 2.6. Implications 28 2.7. Acknowledgments 29 2.8. References 29 Appendix 2.1 Length of conodonts (mm) 32 Appendix 2.2 Height of conodonts (mm) 38 Appendix 2.3 Lateral Area of conodonts (LA≈L*H, mm2) 45 CHAPTER 3 52 EARLY TRIASSIC CONODONTS AND CARBONATE CARBON ISOTOPE RECORD OF THE IDRIJA-ŽIRI AREA, SLOVENIA Abstract 52 3.1. Introduction 53 3.2. Geological settings 54 3.3. Materials and methods 57 3.3.1. Conodont sampling and extraction 57 3.3.2. Unitary Association analysis 67 3.3.3. Carbonate carbon isotope 67 3.4. Conodont biostratigraphy 68 3.4.1. Eurygnathodus costatus Zone (UA 1) 68 3.4.2. Eurygnathodus hamadai Zone (UA 2) 68 3.4.3. Foliella gardenae Zone (UA 3) 69 3.4.4. Neospathodus planus-Neospathodus robustus Zone (UA 4) 69 3.4.5. Platyvillosus corniger Zone (UA 5) 70 3.4.6. Platyvillosus regularis Zone (UA 6) 70 3.4.7. Triassospathodus hungaricus Zone (UA 7) 70 3.4.8. Triassospathodus symmetricus Zone (UA 8) 70 3.4.9. Neospathodus robustispinus Zone (UA 9) 71 3.5. Carbonate carbon isotopes 71 3.6. Discussion 71 3.6.1. The Substage Boundaries 71 3.6.1.1. The Dienerian/Smithian boundary 71 3.6.1.2. The Smithian/Spathian boundary 73 3.6.2. Conodont paleoecology 73 3.6.3. Extinction, recovery and conodonts size 76 3.7. Conclusions 76 3.8. Taxonomic notes (Chen Y.L. & Kolar-Jurkovšek T.) 77 3.9. Acknowledgments 82 3.10. References 82 Appendix 3.1 Data matrix for Unitary Association (UA) analysis 90 Appendix 3.2 Result of carbonate carbon isotope measurements 93 CHAPTER 4 95 GEOMETRIC MORPHOMETRIC ANALYSIS REVEALS THE ALLOMETRY OF ANISIAN (MIDDLE TRIASSIC) SEGMINIPLANATE CONODONTS Abstract 95 4.1. Introduction 95 4.2. Material and geological background 98 4.3. Methods 98 4.4. Results 102 4.4.1. Length measurements 102 4.4.2. Morphometric analysis of the upper view 103 4.4.3. Morphometric analysis of the lateral view 104 4.5 Discussions 106 4.5.1. Size of conodonts 106 4.5.2. Allometry and implications for conodont taxonomy 106 4.6. Conclusions 107 4.7. References 108 CHAPTER 5 113 A REVIEW OF THE EVOLUTION, BIOSTRATIGRAPHY, PROVINCIALISM AND DIVERSITY OF MIDDLE AND EARLY LATE TRIASSIC CONODONTS Abstract 113 5.1. Introduction 114 5.2. Age constraining of conodonts 115 5.3. Lineages and stratigraphic distributions 117 5.3.1. Method 117 5.3.2. Subfamily Novispathodinae Orchard 2005 120 5.3.2.1. Genus Triassospathodus Kozur, 1998 120 5.3.2.2. Genus Nicoraella Kozur 1980 121 5.3.3. Subfamily Pseudofurnishiinae Ramovs, 1977 122 5.3.3.1. Genus Budurovignathus Kozur, 1989 122 5.3.3.2. Genus nov. sp. nov. 123 5.3.3.3. Genus Pseudofurnishius van den Boogaard, 1966 124 5.3.3.4. Genus Mosherella Kozur 1972 125 5.3.3.5. Genus Neocavitella Sudar and Budurov, 1979 126 5.3.3.6. Genus Misikella Kozur and Mock 1974 127 5.3.4. Subfamily Paragondolellinae Orchard 2005 127 5.3.4.1. Genus Paragondolella Mosher 1968 127 5.3.4.2. Genus Quadralella Orchard, 2013 130 5.3.4.3. Genus Kraussodontus Orchard, 2013 132 5.3.4.4. Genus Mazzaella Kilic, Plasencia, Ishida and Hirsch, 2015 133 5.3.4.5. Genus Metapolygnathus Hayashi, 1968 133 5.3.4.6. Genus Parapetella Orchard, 2013 134 5.3.4.7. Genus Carnepigondolella Kozur, 2003 134 5.3.5. Subfamily Epigondolellinae Orchard, 2005 134 5.3.5.1. Genus Acuminatella Orchard, 2013 135 5.3.5.2. Genus Primatella Orchard, 2013 135 5.3.6. Subfamily Neogondolellinae Hirsch, 1994 136 5.3.6.1. Genus Chiosella Kozur 1989 136 5.3.6.2. Genus Neogondolella Bender and Stoppel, 1965 137 5.3.6.3. Genus Celsigondolella Kozur 1968 140 5.3.6.4. Genus Gondolatus Rafek 1977 141 5. 3.7. Subfamily Cornudininae Orchard, 2005 141 5.3.7.1 Genus Cornudina Hirschmann 1959 141 5.3.8. Subfamily Gladigondolellinae Hirsch, 1994 143 5.3.8.1. Genus Gladigondolella Müller, 1962 143 5.3.9. Subfamily uncertain 144 5.3.9.1. Genus Neostrachanognathus Koike, 1998 144 5.4. Conodont provincialism 144 5.5. Conodont diversity 145 5.5.1. The first cycle 147 5.5.2. The second cycle 147 5.6. Conclusions 148 5.7. Acknowledgements 149 5.8. References 149 Appendix 5.1 List conodont holotypes illustrated in this paper 166 Appendix 5.2 Statistic analyses of early, middle and early late Triassic (Carnian) publications 171 PREFACE This thesis focuses on conodonts from the Early Triassic to early Late Triassic. The PhD project started at September, 2011 and Yanlong Chen was personally supported by the China Scholarship Council for his stay in Austria. The studied material of Jiarong section was obtained by himself during his master’s program, but the conodonts size measurements and the writing of the manuscript were conducted at the University of Graz. Parts of Slovenian conodonts were donated by Tea Kolar-Jurkovšek of the Geological Survey of Slovenia and part was obtained by Yanlong Chen after field sampling and laboratory work. Conodonts of Austria (Öfenbachgraben Section and Mendlingbach Section) were donated by Prof. Leopold Krystyn of Vienna University. The integrated study of these conodonts in the present thesis has a profound meaning for biostratigraphy, paleo-environment, biogeography, taxonomy and evolution of Triassic conodonts. And it is a contribution to both the scientific world and the mining and petroleum industry. The PhD candidate (Yanlong Chen) has performed paleontological, biostratigraphical and morphological analyses focusing on the Triassic conodonts. The present thesis is a summary of the work during his doctoral program. Chapter 1 provides a summary and introduction of the paleogeography of the studied areas, the terminology of conodonts shape categories and the state of the art of the research in Triassic conodonts. Chapter 2–5 are papers which have been published, submitted or are ready for submission to a peer reviewed scientific journal. The supplementary information contains the publications following the master thesis in Wuhan (China) and the eight articles co-authored by Yanlong Chen during his PhD programme. Chapter 2 Chen, Y.L., Twitchett, R.J., Jiang, H.S., Richoz, S., Lai, X.L., Yan, C.B., Sun, Y.D., Liu, X.D., Wang, L.N., 2013. Size variation of conodonts during the Smithian- Spathian (Early Triassic) global warming event. Geology, 41(8), 823–826. Chapter 3 Chen, Y.L., Kolar-Jurkovšek, T., Jurkovšek, B., Aljinović, D., Richoz, S. Early Triassic conodonts and carbonate carbon isotope record of Idrija–Žiri area, Slovenia. (Submitted to Palaeogeography Palaeoclimatology Palaeoecology) Chapter 4 Chen, Y.L., Neubauer T.A., Krystyn L., Richoz S. Geometric morphometric analysis reveals the allometry of Anisian (Middle Triassic) segminiplanate conodonts. (Ready for Submission) Chapter 5 Chen, Y.L., Krystyn, L., Orchard, M.J., Lai, X.L., Richoz, S. A review of the evolution, biostratigraphy, provincialism and diversity of Middle and early Late Triassic conodonts. (Papers in Palaeontology, in press) 1 ACKNOWLEDGMENTS I would like to express my thanks to my supervisors, people who have contributed to the thesis, colleagues, and my family. I would like to express here my thankfulness to my supervisors Dr. Sylvain Richoz and Prof. Dr. Werner E. Piller for their guidance, discussion, critical comments and reviews during my PhD program. I would like to thank the Institute of Earth Sciences, University of Graz through Prof. Steffen Birk for providing me chances for attending conferences, summer schools and workshops. Thanks also due to the Institute of Earth Sciences, University of Graz through Prof. Mag. Dr. Walter Kurz for providing me formic acid for dissolving rock samples. I want to thank the staff in the Institute of Earth Sciences, University of Graz, including Thomas J. Suttner, Erika Kido, Markus Reuter, Marco Caporaletti, Evelyn Friesenbichler, Gertraud Bauer, Elisabeth Gülli, Bernhard Hubmann, Magdalena Mandl, Aberra Mogessie, Kurt Stüwe, Gerald Auer, Angela Garcia Gallardo, Patrick Grunert, Juliane Meyer, Claudia Wrozyna, Cyril Mayaud, Jürgen Neubauer, Georg Stegmüller and Franz Tscherne, for their assistance, comment or discussion on my study and/or assistance for my daily life here in Graz. I deeply appreciate my co-workers who spend their precious time to contribute to the present work: Tea Kolar-Jurkovšek, Bogdan Jurkovšek, Dunja Aljinović, Richard J. Twitchett, Xulong Lai, Thomas A. Neubauer, Leopold Krystyn, Mike J. Orchard (GSC, Vancouver), Haishui Jiang, Chunbo Yan, Yadong Sun, Xiaodan Liu and Lina Wang. And I would like to thank Leopold Krystyn and Mike J. Orchard for their reviews and comments on my manuscripts. Finally, I want to express my thankfulness to my wife Lu Zhao, for her strong support for me to do a PhD Program in Europe, to my son Junwen Chen for bringing happiness to our family.