La ilustración de la portada de esta memoria es creación original del Dr. Carlos de Miguel. La ilustración de los fondos del título de cada capítulo es creación original de la autora de esta memoria, Aitziber Suárez. La autora de esta tesis doctoral ha sido financiada por una beca de Formación de Personal Investigador no doctor del Programa Predoctoral del Gobierno Vasco. UNIVERSIDAD DEL PAÍS VASCO (UPV-EHU) FACULTAD DE CIENCIA Y TECNOLOGÍA DEPARTAMENTO DE ESTRATIGRAFÍA Y PALEONTOLOGÍA Late Pleistocene vertebrate assemblages from Artazu VII and Artazu VIII sites (Arrasate, northern Iberian Peninsula): Palaeobiology and Palaeoecology TESIS DOCTORAL Aitziber Suárez Bilbao Bajo la dirección de los Drs.: Xabier Murelaga Bereicua Naroa Garcia Ibaibarriaga Departamento de Estratigrafía y Departamento de Geografía, Paleontología Prehistoria y Arqueología Facultad de Ciencia y Tecnología Facultad de Letras Universidad del País Vasco Universidad del País Vasco (UPV-EHU) (UPV-EHU) Leioa, octubre de 2018 (c)2019 AITZIBER SUAREZ BILBAO “No se trata de qué fuerte golpees, se trata de cuanto eres capaz de soportar ser golpeado y seguir moviéndote. Así es como se gana” Sylvester Stallone “Cada día sabemos más y entendemos menos” Albert Einstein GENERAL INDEX GENERAL INDEX ACKNOWLEDGEMENTS ................................................................................ 13 ABSTRACT/LABURPENA/RESUMEN EXTENDIDO ..................................... 21 1. INTRODUCTION .......................................................................................... 29 1.1. The current climate in the Iberian Peninsula and Cantabrian region .... 31 1.2. Climatic cycles during the Late Pleistocene .......................................... 33 1.3. Late Pleistocene palaeontological sites in the Iberian Peninsula .......... 37 1.4. Organization of thesis ............................................................................ 38 1.5. Principal goals ....................................................................................... 40 2. LOCATION OF SITES................................................................................... 41 2.1. Geographical settings ............................................................................ 43 2.2. Geological context ................................................................................. 44 2.3. Description of the cavities ...................................................................... 46 2.3.1. Artazu VII cavity .................................................................... 46 2.3.2. Artazu VIII cavity ................................................................... 47 3. MATERIALS AND METHODS ..................................................................... 51 3.1. Field techniques: excavation works and sample collection ................... 53 3.1.1. Excavation works and palaeontological samples ................. 53 A. Artazu VII .................................................................................... 56 Small vertebrates ........................................................................ 57 Large mammals .......................................................................... 57 Avifauna ...................................................................................... 57 B. Artazu VIII .................................................................................. 58 Small vertebrates ........................................................................ 59 Large mammals .......................................................................... 59 Avifauna ...................................................................................... 60 Pollen .......................................................................................... 60 5 3.1.2. Sedimentological and mineralogical samples ...................... 62 3.1.3. Micromorphology samples .................................................... 62 3.2. Laboratory techniques ........................................................................... 63 3.2.1. Dating techniques ................................................................. 63 Radiocarbon Dating (AMS) ........................................ 63 Amino Acid Racemisation (AAR) ............................... 63 A. Artazu VII .................................................................................... 64 B. Artazu VIII ................................................................................... 64 Uranium/Thorium (U/Th) ............................................ 65 3.2.2. Palaeontological samples ..................................................... 66 Small vertebrates .................................................................. 67 Identification of the material ....................................... 70 Biometry of Soricidae lower dental elements ............. 78 Quantification of the material ..................................... 79 Specific diversity analysis ........................................... 79 Habitat assemblage .................................................... 80 - Type of habitats ................................................ 80 - Factors to be taken into consideration ............... 80 Large mammals ................................................................... 81 Collection number ...................................................... 81 Identification of the material ....................................... 82 Quantification of the material ..................................... 83 Determination of sex ................................................... 83 Determination of the age at death ............................... 83 Osteometric measurements ........................................ 83 Avifauna ............................................................................... 84 Identification of the material ....................................... 84 Quantification of the material and Osteometric measurements .................................................................... 86 Palaeopalinology ................................................................. 86 3.2.3. Sedimentological and mineralogical samples ...................... 87 Granulometric analyses ....................................................... 87 6 GENERAL INDEX Mineralogical analyses (XRD) ............................................. 87 Micromorphology samples.................................................... 87 3.2.4. Geochemical samples .......................................................... 88 4. TAPHONOMIC REMARKS ........................................................................... 91 4.1. Small vertebrates .................................................................................... 93 4.2. Large mammals ..................................................................................... 95 4.3. Avifauna ................................................................................................. 96 5. SYSTEMATIC PALAEONTOLOGY ............................................................ 101 5.1. Mammals .............................................................................................. 103 Arvicola amphibius ...................................................................... 103 Arvicola sapidus .......................................................................... 105 Microtus (Microtus) agrestis ........................................................ 106 Microtus (Microtus) arvalis .......................................................... 106 Microtus (Terricola) sp. ............................................................... 107 Microtus (Alexandromys) oeconomus ......................................... 108 Chionomys nivalis ....................................................................... 109 Pliomys lenki ............................................................................... 109 Apodemus sylvaticus-flavicollis ................................................... 110 Erinaceus europaeus .................................................................. 112 Sorex (Sorex) araneus-coronatus ............................................... 112 Sorex (Sorex) minutus ................................................................ 114 Neomys sp. ................................................................................. 114 Talpa sp. ..................................................................................... 116 Felis silvestris .............................................................................. 117 Lynx sp. ...................................................................................... 117 Panthera pardus .......................................................................... 118 Panthera leo spelaea .................................................................. 120 Canis lupus .................................................................................. 121 Cuon alpinus ............................................................................... 121 Vulpes vulpes .............................................................................. 122 7 Ursus spelaeus............................................................................ 123 Martes martes ............................................................................. 124 Meles meles ................................................................................ 124 Mustela nivalis ............................................................................. 125 Mustela putorius .........................................................................