Departament de Biologia Animal, de Biologia Vegetal i d’Ecologia Unitat d’Antropologia Biològica Miocene carnivorans from the Vallès-Penedès Basin (NE Iberian Peninsula) Josep Maria Robles Giménez Tesi Doctoral 2014 A mi padre y familia. INDEX Index .......................................................................................................................... 7 Preface and Acknowledgments [in Spanish] ....................................................... 13 I.–Introduction and Methodology ........................................................................ 19 Chapter 1. General introduction and aims of this dissertation .......................... 21 1.1. Aims and structure of this work .............................................................. 21 Motivation of this dissertation ................................................................ 21 Type of dissertation and general overview ............................................. 22 1.2. An introduction to the Carnivora ............................................................ 24 What is a carnivoran? ............................................................................. 24 Biology .................................................................................................... 25 Systematics and phylogeny ...................................................................... 28 Evolutionary history ................................................................................ 42 1.3. Carnivoran anatomy ................................................................................ 45 Dentition .................................................................................................. 45 Skull (cranium and mandible) ................................................................. 49 Postcranium ............................................................................................ 51 1.4. Previous research on fossil carnivorans from the Vallès-Penedès Basin ........................................................................................................ 53 Early studies ............................................................................................ 53 Modern studies ........................................................................................ 56 State of the art ......................................................................................... 59 Chapter 2. Materials and methods ..................................................................... 61 2.1. Sample, anatomical nomenclature and measurements ............................ 61 Studied sample ......................................................................................... 61 Anatomical nomenclature ....................................................................... 62 Measurements .......................................................................................... 62 2.2. Analytical methods .................................................................................. 63 Taxonomy and comparative anatomy ..................................................... 63 Phylogenetic reconstruction .................................................................... 63 Computed tomography ............................................................................ 65 Chapter 3. Geology of the Vallès-Penedès Basin .............................................. 67 8 JOSEP M. ROBLES Miocene carnivorans from the Vallès-Penedès Basin (NE Iberian Peninsula) 3.1. An introduction to the Vallès-Penedès Basin ...........................................67 Geographic situation ................................................................................67 Tectonosedimentary history .....................................................................67 Main stratigraphic units of the Vallès-Penedès Basin. ............................69 3.2. The vertebrate fossil record from the Vallès-Penedès Basin ...................75 II.–Results ................................................................................................................83 Chapter 4. Updated chronology for the Miocene hominoid radiation in Western Eurasia ..............................................................................................85 4.1. Abstract ....................................................................................................87 4.2. Introduction ..............................................................................................87 4.3. Results and discussion .............................................................................87 Oldest Eurasian hominoid? .....................................................................87 Middle Miocene sites ...............................................................................87 Late Miocene sites ....................................................................................89 Latest hominoids from Western Eurasia ..................................................90 4.4. Conclusions ..............................................................................................90 4.5. Materials and methods .............................................................................91 Chronological framework: Regional timescales and correlation ............91 4.6. Acknowledgments ....................................................................................91 4.7. Literature cited .........................................................................................91 4.8. Supporting Information Appendix ...........................................................93 Systematics of Western Eurasian hominoids ...........................................93 Regional chronological systems .............................................................101 The Miocene record of the Vallès-Penedès Basin: biostratigraphy and magnetostratigraphy ..................................................................102 References ..............................................................................................106 4.9. Supporting Information Dataset .............................................................110 How to use this database .......................................................................113 Locality database ...................................................................................115 References ..............................................................................................137 Chapter 5. New craniodental remains of Trocharion albanense Major, 1903 (Carnivora, Mustelidae), from the Vallès-Penedès Basin (Middle to Late Miocene, Barcelona, Spain) ..........................................................................141 5.1. Abstract ..................................................................................................143 INDEX 9 5.2. Introduction ........................................................................................... 143 The genus Trocharion in the Iberian Peninsula .................................... 143 Historical background ........................................................................... 143 5.3. Stratigraphy and age .............................................................................. 145 5.4. Materials and methods .......................................................................... 145 Nomenclature ........................................................................................ 145 Abbreviations ........................................................................................ 145 Studied material .................................................................................... 145 Comparative sample .............................................................................. 146 Cladistic analysis .................................................................................. 146 5.2. Systematic paleontology ....................................................................... 146 Description .......................................................................................... 150 5.3. Cladistic analysis ................................................................................... 153 5.4. Discussion ............................................................................................. 154 Comparison with other Middle Miocene European musteloids ......... 154 Nomenclatural and taxonomic issues ................................................. 154 Systematics and phylogeny ................................................................. 155 Zoogeography remarks ....................................................................... 155 5.5. Conclusions ........................................................................................... 156 5.6. Acknowledgments ................................................................................. 156 5.7. Literature cited ...................................................................................... 156 Chapter 6. New craniodental remains of the barbourofelid Albanosmilus jourdani (Filhol, 1883) from the Miocene of the Vallès-Penedès Basin (NE Iberian Peninsula) and the phylogeny of the Barbourofelini ................ 159 6.1. Abstract ................................................................................................. 161 6.2. Introduction ........................................................................................... 161 Sansanosmilus and Barbourofelis ......................................................