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Oological Record of Dinosaurs in South-Central Pyrenees (SW Europe): Parataxonomy, diversity and biostratigraphical implications Albert Garcia Sellés ADVERTIMENT. La consulta d’aquesta tesi queda condicionada a l’acceptació de les següents condicions d'ús: La difusió d’aquesta tesi per mitjà del servei TDX (www.tdx.cat) ha estat autoritzada pels titulars dels drets de propietat intel·lectual únicament per a usos privats emmarcats en activitats d’investigació i docència. No s’autoritza la seva reproducció amb finalitats de lucre ni la seva difusió i posada a disposició des d’un lloc aliè al servei TDX. No s’autoritza la presentació del seu contingut en una finestra o marc aliè a TDX (framing). Aquesta reserva de drets afecta tant al resum de presentació de la tesi com als seus continguts. En la utilització o cita de parts de la tesi és obligat indicar el nom de la persona autora. ADVERTENCIA. 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OOLOGICAL RECORD OF DINOSAURS IN SOUTH-CENTRAL PYRENEES (SW EUROPE): PARATAXONOMY, DIVERSITY AND BIOSTRATIGRAPHICAL IMPLICATIONS Albert Garcia Sellés Universitat de Barcelona Departament d’Estratigrafia, Paleontologia i Geociències Marines 2012 OOLOGICAL RECORD OF DINOSAURS IN SOUTH-CENTRAL PYRENEES (SW EUROPE): PARATAXONOMY, DIVERSITY AND BIOSTRATIGRAPHICAL IMPLICATIONS Albert Garcia Sellés Memòria de Tesi Doctoral Universitat de Barcelona Departament d’Estratigra a, Paleontologia i Geociències Marines 2012 Front page: Artwork based on ilustration of Albert García Ames (Daltonico Studio) OOLOGICAL RECORD OF DINOSAURS IN SOUTH-CENTRAL PYRENEES (SW EUROPE): PARATAXONOMY, DIVERSITY AND BIOSTRATIGRAPHICAL IMPLICATIONS Memòria presentada per Albert Garcia Sellés per optar al Grau de Doctor en Ciències de la Terra per la Universitat de Barcelona, sota la direcció del Dr. Àngel Galobart Lorente, Cap del Grup de Recerca del Mesozoic del Institut Català de Paleontologia Miquel Crusafont amb la tutela del Dr. Xavier Dalclòs Martínez, Professor titular del Departament d’Estratigra a, Paleontologia i Geociències Marines de la Universitat de Barcelona Als meus pares; “...Lightly on the knee she settles, " nds a nesting-place be tting, where to lay her eggs in safety. here she builds her humble dwelling, lays her eggs within, at pleasure, six, the golden eggs she lays there, then a seventh, an egg of iron; sits upon her eggs to hatch them... ...Quick the maiden moves her shoulders, shakes her members in succession, shakes the nest from its foundation, and the eggs fall into ocean, dash in pieces on the bottom... ...From one half the egg, the lower, grows the nether vault of Terra: from the upper half remaining, grows the upper vault of Heaven; from the white part come the moonbeams, from the yellow part the sunshine, from the motley part the starlight, from the dark part grows the cloudage...” The Kalevala. Rune I - Birth of Wainamoinen Translated by John Martin Crawford (1888) CONTENT Acknowledgements (in Catalan) 11 Resume (in Catalan) 15 Prologue Motivation, Problematic and Objectives 29 Section 1 Introduction and Contextualization 35 Chapter 1 Historical overview on the study of dinosaur eggs: special attention to South-Central Pyrenees 37 record and problematic Chapter 2 Geographical and Geological settings 41 2.1. Geographical settings 41 2.2. Geological settings 41 2.2.1. Uppermost Cretaceous continental deposits in South-Pyrenean zone: Generalities 42 2.2.2. Àger Syncline 44 2.2.3. Tremp Basin 45 2.2.4. Coll de Nargó Syncline 47 2.2.5. Vallcebre Syncline 48 Chapter 3 Material, Methods and Parataxonomy 51 3.1. Material and Methods 51 3.1.1. Acquisition of studied material 51 3.1.1.1. Sampling in Coll de Nargó (Alt Urgell) 51 3.1.1.2. Sampling in Àger and Tremp basins (Noguera and Pallars Jussà) 52 3.1.2. Preparation of fossil eggs and eggshells 53 3.1.3. Scanning Electron Microscopy (SEM) 53 3.1.4. Thin-section preparation 54 3.1.5. Descriptions 56 7 3.2. Introduction to parataxonomycal classi cation of fossil eggs 57 3.2.1. Eggshell structure 57 3.2.2. Terminology 59 3.2.3. Basic types and morphotypes of eggshell structures 65 Section 2 Systematic Paleontology and Phylogeny of fossil eggs 69 Chapter 4 Record and Classi cation of sauropod eggs 71 4.1. Sauropodomorph egg record: an overview on its biogeographical time distribution 71 4.1.1. Sauropodomorph egg record 73 4.1.1.1. Triassic 73 4.1.1.2. Jurassic 75 4.1.1.3. Cretaceous 76 4.1.1.3.1. Early Cretaceous 76 4.1.1.3.2. Late Cretaceous 78 4.1.2. Fossil egg record and paleogeographical history of sauropodomorph dinosaurs 84 4.1.2.1. The origin of sauropodomorph (Late Triassic to Early Jurassic) 84 4.1.2.2. Rise of sauropods and the domain of neosauropoda (Middle Jurassic to Late Jurassic) 85 4.1.2.3. European “land bridge”: the dispersion of macronaria sauropods and the rise of titanosaurs 86 (Early Cretaceous) 4.1.2.4. Sauropod “mid-Cretacious hatus” in northern continents and new ootaxa (Middle cretaceous: 87 from Aptian to Santonian) 4.1.2.5. Austral immigration of titanosaurs (Late Cretaceous: from Campanian to Maastrichtian) 88 4.2. Problematics in megaloolithid eggs: diagnostic criteria and validity of oospecies 91 4.2.1. Structural levels and characters 92 4.2.2. The Megaloolithid Groups (MG’s) 97 4.2.2.1. Megaloolithid Group 1 (MG1) 97 4.2.2.2. Megaloolithid Group 2 (MG2) 100 4.2.2.3. Megaloolithid Group 3 (MG3) 103 4.2.2.4. Ungrouped megaloolithid oospecies 111 4.3 Megaloolithid eggs in the Upper Cretaceous of Southern Pyrenees from Catalonia 113 8 Chapter 5 Non-sauropod eggshells from the Upper Cretaceous of Southern Pyrenees 119 5.1. Cairanoolithid eggs from the Upper Cretaceous of Southern Pyrenees 119 5.1.1. Introduction 119 5.1.2. Systematic Paleontology 120 5.1.3. Discussion 123 5.2. The “thin eggshells” from the Late Cretaceous in Southern Pyrenees 126 5.2.1. Introduction 126 5.2.2. Systematic Paleontology 126 Chapter 6 Phylogenetyc analysis on dinosaur fossil eggs 137 6.1. Phylogenetical relationship between megaloolithid oospecies 137 6.1.1. Results of the phylogenetic analysis 139 6.1.2. Discussion of phylogenetic analysis 141 6.2. Phylogenetical analysis of Cairanoolithus 142 6.2.1. Results of the phylogenetic analysis 143 6.1.2. Discussion of phylogenetic analysis 145 6.2.3. Considering the taxonomical af nity of Cairanoolithus 145 Section 3 Oological diversity, temporal variability and taxonomical relationships 147 Chapter 7 Oological diversity in Southern Pyrenees 149 7.1. Comparison with other sites 151 7.2. Temporal distribution in South Pyrenees: implications for taxonomical diversity, nesting site preference and 152 environmental factors 7.3. Possible taxonomical relationship between megaloolithid eggs and sauropod fauna in the Upper Cretaceous 156 of Europe. Section 4 Biostratigraphical implications 161 Chapter 8 Bioestatigraphy of the Upper Cretaceous of Southern Europe based on the Megaloolithus 163 oospecies succession 8.1. Introduction 163 9 8.2. Biostratigraphy at Pinyes nesting area (Coll de Nargó Syncline) 164 8.3. Biostratigraphy of the Àger and Tremp basins 167 8.4. Considerations about the biostratigraphy in the Vallcebre Syncline 168 8.5. Updating the European megaloolithid biochronology 168 General conclusions 175 References 183 Appendices 217 Appendix I 219 Appendix II 229 Apendix III 231 10 Acknowledgements (in Catalan) Sens dubte aquesta és la part més esperada i llegida de qualsevol tesi doctoral, on tothom espera veure-s’hi re exat o on trobar-hi anècdotes divertides. Però el protocol marca les normes, així que començarem per la part formal: La present tesi ha esta possible gràcies a la contribució, suport i ajuda rebuda de nombrosos companys, amics, col·legues i familiars. A tots ells els hi vull transmetre el meu més profund agraïment. L’autor de la tesi a estat bene ciari d’una beca predoctoral 2009 FI 09 atorgada per l’Agència de Gestió d’Ajuts Universitaris i de Recerca (AGAUR) de la Generalitat de Catalunya. Els treballs realitzats en el marc d’aquesta tesi han estat nançats pels següents projectes de la Subdirección General de Proyectos de Investigación del Ministerio de Ciencia e Innovación i del Ministerio de Economia y Competitividad. Aquests projectes són: “Evolucion de los dinosaurios en el este iberico, y su entorno, durante el cretacico: sistematica e inferencias paleobiologicas y paleoecologicas” project number CGL2008-06533-c03-01/BTE i “El " n de una era: la extinción de los dinosaurios, una prespectiva europea” project number GL2011-30069-c02-01/ BTE, ambdos sota la direcció del Dr.