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TESIS CORRAL ARROYO JOSE CARMELO.Pdf Upper Cretaceous neoselachians from the Basque-Cantabrian Region (Northern Spain) “Shark teeth are the most commonly collected vertebrata fossil. They appear on beaches, prairies, mountaintops, and deserts, as well as in riverbeds. They have also been found in the Antarctic and the deepest part of the ocean. In the living shark, teeth are constantly produced and shed; a typical carcharhinid, such as the lemon shark Negaprion brevirostris, may produce 20,000 teeth in its first 25 years, and may live as long as 50 years.” Gordon Hubbell, 1996 DOCTORAL RESEARCH THESIS UPPER CRETACEOUS NEOSELACHIANS from the Basque-Cantabrian Region (Northern Spain) Thesis submitted by José Carmelo Corral Arroyo for the degree of Doctor in Geology within the Department of Stratigraphy and Palaeontology Universidad del País Vasco / Euskal Herriko Unibertsitatea / University of the Basque Country 2018 (c)2018JOSE CARMELO CORRAL ARROYO Thesis directors Dr Javier Murelaga Bereicua | Dr Xabier Pereda-Suberbiola iv ‘Ad Divisionem piscium priorem pertinent illi dentes Squalorum qui Glossopetrae vulgo appellantur, & qui in nostra quoque formatione cretacea, varii locis & in variis stratis dispersi jacent.’ Sven Nilsson, 1827 (Petrificata Suecana formationis cretaceae, descripta et iconibus illustrata. Pars prior, Vertebrata et Mollusca) This thesis is dedicated to my late father, Ernesto, who fostered mi interest in fossils at an early age, and to my always encouraging mother, Evangelina. Thanks for all your support CONTENTS Purpose of investigation and objectives xxi Outline of the thesis and Statement on the contribution of others xxiii Acknowledgements xxvii Abstract xxix Resumen extendido xxxi INTRODUCTION Chapter 1 Historical, geographic, and geological settings 1.1 Historical review of fossil neoselachian research in the Basque-Cantabrian Region 3 1.2 Geography of the study area 9 Southwestern region – Plateau of La Lora de la Pata del Cid (Burgos) 9 Central region – Quintanilla Ojada (Burgos) 12 Eastern region (Álava and Treviño County in Burgos) 12 1.3 A brief geological overview of the Basque-Cantabrian Basin 16 Boundaries and structural organization 16 Sedimentary basin fill with particular attention to the study area 18 Chapter 2 Material and methods 2.1 Introduction 33 2.2 General field techniques (specimen collection) 36 2.3 Screen washing 39 2.4 Sorting and picking 41 2.5 Preparation of palaeontological remains 43 Mechanical preparation 47 Chemical preparation techniques 50 Consolidation 54 2.6 Geochemistry methods 55 2.7 Analytical methods for palaeobiogeographical reconstruction 55 2.8 Photography and Illustrations 57 Whitening specimens 59 Infographics and drawings 60 2.9 Management of the elasmobranch collection 61 General considerations 61 xv Conservation of the specimens discussed in this thesis 61 2.10 Special case: the preparation of a mosasaurid vertebra from Jauregi (Álava) 62 Chapter 3 An overview of living and fossil elasmobranchs 3.1 Introduction – What is an elasmobranch? 67 Classification of living elasmobranchs 67 A general overview of the Elasmobranchii fossil record 72 3.2 Endoskeleton 75 Cartilaginous skeleton 77 3.3 Exoskeleton 78 Tooth morphogenesis 78 Tooth histology and composition 80 Dental characters 84 Tooth diversity and orientation 84 Other exoskeletal structures (placoid scales, thorns, tail spines) 89 3.4 Teeth patterns and specialised trophic adaptations 90 3.5 Nomenclature and systematics of elasmobranchs 92 Box 3.1 Most common dental terms (English–Spanish) used to describe shark teeth 96 GEOLOGY AND PALAEONTOLOGY OF THE STUDIED AREAS Chapter 4 Stratigraphy and palaeontology of the Barrio Panizares beds (Nidáguila Formation, Coniacian) from Northern Spain 4.1 Location and geological setting 101 4.2 Stratigraphy 101 Facies interpretation 103 Stratigraphical position 105 Chapter 5 Stratigraphy and palaeontology of the upper Campanian Gometxa beds (Vitoria- Gasteiz, Álava) 5.1 Introduction 111 5.2 Location and geological setting 112 5.3 Stratigraphy 116 Review of the Gomecha Member (GM) 116 5.4 Some general taphonomic features observed on the elasmobranch teeth 128 Chapter 6 Stratigraphy and palaeontology of the upper Campanian Vitoria Pass beds (Vitoria- Gasteiz, Álava) 6.1 Introduction 129 6.2 Location and geological setting 130 6.3 Stratigraphy 132 Stratigraphical description 132 Stratigraphical interpretation 138 6.4 Review of the Montes de Vitoria Formation (MVF) 142 xvi Eguileta Member (EM) 143 6.5 Biostratigraphy and age assigned to the Vitoria Pass site 145 The planktonic foraminiferal zones 145 Ammonite zonal scheme 147 Age assigned to the Vitoria Pass beds 151 Chapter 7 Geology and geochemistry of the Maastrichtian Quintanilla la Ojada beds (Valle de Losa, Burgos) 7.1 Introduction 153 7.2 Location and geological setting 153 7.3 Stratigraphy 156 Stratigraphical description 156 Stratigraphical interpretation 158 7.4 Geochemistry and Taphonomy 159 Rare Earth Elements analysis 159 Taphonomy 161 7.5 Age of the deposits 164 Chapter 8 Stratigraphy and palaeontology of the upper Maastrichtian Albaina beds (Treviño County, Burgos) 8.1 Introduction 167 8.2 Location and geological setting 168 8.3 Stratigraphy 171 Sobrepeña Formation (SPF) 171 Torme Formation (TF) 177 8.4 Taphonomic features of the selachian deposit 191 Chapter 9 Stratigraphy and palaeontology of the upper Maastrichtian Entzia beds (Urbasa, Álava) 9.1 Introduction 193 9.2 Location and geological setting 193 9.3 Stratigraphy 194 Puerto de Olazagutía Formation (POF) 194 9.4 Ammonite biostratigraphy 201 NEOSELACHIAN (SHARKS AND RAYS) SYSTEMATICS Chapter 10 A rostral spine of the sawfishOnchosaurus (Neoselachii, Sclerorhynchidae) from the Barrio Panizares beds (Northern Spain), with a revision of the genus 10.1 Introduction 207 10.2 Systematic palaeontology 207 10.3 Redescriptions of the type material of Onchosaurus 212 Onchosaurus radicalis Gervais, 1852 212 Onchosaurus pharao (Dames, 1887a) 212 xvii Chapter 11 Neoselachians from the Gometxa site (Gomecha Member, upper Campanian) 11.1 Introduction 215 11.2 Systematic palaeontology 215 Chapter 12 Neoselachians from the Vitoria Pass site (Eguileta Member, upper Campanian) 12.1 Introduction 257 12.2 Systematic palaeontology 257 Chapter 13 Neoselachians from the Quintanilla la Ojada site (Valdenoceda Formation, Maastrichtian) 13.1 Introduction 279 13.2 Systematic palaeontology 279 Chapter 14 Neoselachians from the Albaina site (Torme Formation, upper Maastrichtian) 14.1 Introduction 303 14.2 Systematic palaeontology 303 Chapter 15 Neoselachians from the Entzia sites (Puerto de Olazagutía Formation, upper Maastrichtian) 15.1 Introduction 339 15.2 Systematic palaeontology 339 RESEARCH ON THE NEOSELACHIAN PALAEOECOLOGY AND PALAEOBIOGEOGRAPHY Chapter 16 Stratigraphical and palaeobiogeographical distribution of the sawfishOnchosaurus (Neoselachii, Sclerorhynchidae) 16.1 Introduction 349 16.2 Distribution of Onchosaurus 349 Stratigraphical distribution 349 Palaeobiogeographical distribution 350 Chapter 17 Palaeoecology and palaeobiogeography of the Campanian selachians in the Gometxa and Vitoria Pass beds 17.1 Introduction 355 17.2 Species diversity and faunal affinities 356 The Gometxa site 356 The Vitoria Pass site 358 17.3 Palaeoecology and biogeographical faunal patterns 358 17.4 Global patterns in the biogeographical distribution of neoselachians during the Campanian time 366 Common multivariate ordinations and parsimony analysis 366 Discussion of the results 370 xviii Chapter 18 Bite marks attributed to a shark on a mosasaurid vertebra (Vitoria Sub-basin, Campanian) 18.1 Introduction 385 18.2 Location and geological setting 386 18.3 Description of the specimen 388 18.4 Discussion 392 Chapter 19 Marine vertebrate predators (Actinopterygii, Chondrichthyes, and Mosasauridae) in the Campanian of the Basque-Cantabrian Basin 19.1 Introduction 397 19.2 Marine vertebrate groups 398 Actinopterygii (Bony Fish) 398 Selachians (Chondrichthyes: Euselachii: Neoselachii) 398 Mosasaurids (Squamata: Mosasauridae) 398 19.3 Selachian ecomorphotypes 401 Chapter 20 Selachian palaeoecology, stratigraphical distribution, and palaeobiogeography of the Maastrichtian assemblages (Quintanilla la Ojada, Albaina, and Entzia) 20.1 Introduction 403 20.2 Palaeoecology 404 20.3 Species diversity, faunal affinities, and palaeobiogeographical implications 408 CONCLUSIONS Chapter 21 Conclusions 21.1 General conclusions 417 21.2 Conclusions on the Coniacian stratigraphy and associated faunal remains 418 21.3 Conclusions on the Campanian stratigraphy and associated faunal remains 418 21.4 Conclusions on the cut marks on a mosasaurid vertebra from the Campanian of the Vitoria Sub-basin 421 21.5 Conclusions on the marine vertebrate predators (Actinopterygii, Chondrichthyes, and Mosasauridae) in the Campanian of the Basque-Cantabrian Basin (B-CB) 422 21.6 Conclusions on the Maastrichtian stratigraphy and associated faunal remains 422 References 429 Systematic index 471 xix It all began with ... Some of my first findings of shark’s teeth discovered (circa 1983) in the Campanian marls of the Vitoria Ranges, near the city of Vitoria-Gasteiz. These specimens are now housed in the Ara- bako Natur Zientzien Museoa / Museo de Ciencias Naturales de Álava (Basque Country, Spain). Photography by geologist Agustín Pascual, ca. 1984. | Purpose of investigation and objectives Purpose of investigation and objectives My interest in fossil shark teeth began in 1983 when I stumbled upon several teeth in an abandoned quarry not far from my birth town,
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